protosalvinia dawson and associated ......b. general famennian stratigraphy of the appalachian and...

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J. Paleont., 83(1), 2009, pp. 70–79Copyright � 2009, The Paleontological Society0022-3360/09/0083-70$03.00

PROTOSALVINIA DAWSON AND ASSOCIATED CONODONTS OF THE UPPERTRACHYTERA ZONE, FAMENNIAN, UPPER DEVONIAN, IN THE

EASTERN UNITED STATES

D. JEFFREY OVER1 REMUS LAZAR,2 GORDON C. BAIRD,3 JUERGEN SCHIEBER4 AND FRANK R. ETTENSOHN5

1Department of Geological Sciences, State University of New York College at Geneseo, Geneseo, 14454, �[email protected]�;2Exxonmobil Upstream Research Co., Houston, Texas 77027; 3Department of Geological Sciences, State University of New York College at Fredonia,

Fredonia, 14063; 4Department of Geological Sciences, Indiana University, Bloomington, 47405; 5Department of Earth and Environmental Sciences,University of Kentucky, Lexington, 40506

ABSTRACT—Protosalvinia first occur in association with conodonts of the Upper trachytera Zone and below the Three Lick Bed in the OhioShale and the Ellicott Shale of the central and northern Appalachian Basin, as well as in the Clegg Creek Member of the New Albany Shaleof the Illinois Basin. In the Chattanooga Shale of the southern Appalachian Basin, Protosalvinia are found no lower than the Upper marginiferaZone or associated with obviously reworked conodonts in the Middle expansa Zone. Regionally Protosalvinia are associated with a disconformityand may be found with conodonts of the Lower expansa Zone.

INTRODUCTION

PROTOSALVINIA DAWSON, 1884, also widely known by the ju-nior synonym Foerstia White, 1923, are the remains of an

enigmatic Late Devonian plant-like organism of possible terres-trial origin (Phillips et al., 1972; Gray and Boucot, 1979; Ro-mankiw et al., 1988). The macroscopic organic remains areknown from numerous localities in the eastern United States,largely from offshore marine mudrocks, in a relatively narrowstratigraphic interval (Schopf and Schwietering, 1970; Murphy,1973; Hassenmueller et al., 1983; Conkin, 1985; Ettensohn et al.,1988). Protosalvinia has also been found in North Dakota (Hol-land et al., 1987), Montana (Cross, 1982, in Matthews, 1983),Oklahoma (Hass, 1956), southern Ontario (Winder, 1966) andBrazil (Dawson, 1884; Niklas et al., 1976; Loboziak et al., 1997).Hannibal (1994) noted that the carbonized cephalopod aptychusSidetes Giebel, 1847 may have locally been misidentified as Pro-tosalvinia. The converse is also true (see Winder, 1966), and un-confirmed reports should be considered with caution. Collectionsof Protosalvinia-bearing strata in New York, Pennsylvania, Ohio,Indiana, Kentucky, and Tennessee (see locality register in Appen-dix 1) yielded conodonts that constrain the base of the Protosal-vinia Interval to no higher than the Upper trachytera Zone in theeastern United States. The lower occurrences of Protosalvinia inthe rhomboidea and marginifera zones, as proposed by Sandberget al. (1994) and proliferated by Loboziak et al. (1997), are notsupported by direct conodont evidence. It is not clear if Proto-salvinia associated with conodonts typical of the Lower expansaZone are a higher occurrence or reworked.

STRATIGRAPHY

Protosalvinia in the Appalachian Basin are known from themedial Ellicott Shale Member of the Chadokoin Formation in theupper part of the Conneaut Group in New York and Pennsylvania,from the Chagrin and Huron shales in Ohio, Kentucky, and WestVirginia below the Three Lick Bed of Provo et al. (1978), andfrom the upper Gassaway Member of the Chattanooga Shale inKentucky, Tennessee, and Georgia (Fig. 1). In the Illinois Basinof Indiana and Kentucky, Protosalvinia are found in the lowerClegg Creek Member of the New Albany Shale. The Protosal-vinia-bearing interval is typically associated with strata in thelower part of the middle Huron Shale, just above the high gamma-ray interval of the lower Huron Shale and its equivalents in the

Ohio, Chattanooga, New Albany, and Antrim shales (Hassen-mueller et al., 1983; Ettensohn, 1992). Winder (1966), Matthews(1983), and Sandberg et al. (1994) reported Protosalvinia fromthe Antrim and Kettle Point formations of Michigan, Ontario, andIndiana in the Michigan Basin.

PALEONTOLOGY

Protosalvinia remains range from 1 to 20 mm in width, 5 mmbeing typical, characteristically flattened and often bilobed, witha reticulated cellular pattern (Fig. 2). Complete or fragmentedProtosalvinia are often found in high concentration on beddingplanes in marine mudrocks, but also in lag beds and shell layers.Conodonts in this study were recovered from acid residues ofshell-rich layers, as well as from shale bedding plane surfacescollected from outcrops and drill cores. The conodont zonationused for the Famennian, inclusive of the Protosalvinia Interval,is the scheme proposed by Ziegler (1962a) and most recentlymodified by Ziegler and Sandberg (1990), which is here plottedagainst the Devonian radiometric time scale presented by Kauf-mann (2006; Figure 1). The ranges of conodonts recovered justbelow or within the Protosalvinia Interval include relatively longranging taxa such as Palmatolepis schindewolfi Muller, 1956 andPelekysgnathus inclinatus Thomas, 1949, but also taxa that haveshort ranges (e.g., Palmatolepis marginifera marginifera Helms,1959, Palmatolepis marginifera utahensis Ziegler and Sandberg,1984, P. glabra distorta Branson and Mehl, 1934a, and P. per-lobata grossi Ziegler in Kronberg, Pilger, Scherp, and Ziegler,1960; Fig. 3). The occurrence of Palmatolepis rugosa rugosaBranson and Mehl, 1934a from the New Albany Shale (localityA, Table 1) associated with Palmatolepis falcata (Helms, 1959),P. perlobata grossi, P. perlobata sigmoidea Ziegler, 1962b, P.glabra distorta, and Polylophodonta confluens (Ulrich and Bas-sler, 1926), which are not known to range into the Lower expansaZone, may be a lower occurrence than previously reported or anindicator of the Lower expansa Zone and a reworked fauna ofwell-preserved older conodonts. The occurrence of Palmatolepisfalcata, P. glabra distorta, and P. glabra angusta Capkinoglu,1997 in strata above the basal Protosalvinia-bearing lag horizonthat contain P. rugosa rugosa (Table 1, locality A) indicates thata lowering of the range of P. rugosa rugosa to the Upper trach-ytera Zone is warranted. The absence of conodonts indicative ofhigher zones further supports this assessment.

In the Ellicott Shale Member of the Chadokoin Formation in

71OVER ET AL.—DEVONIAN PROTOSALVINIA (FOERSTIA) OF EASTERN USA

FIGURE 1—A. Distribution of Upper Devonian marine strata in eastern NorthAmerica, group and formation names of predominant Protosalvinia-bearing strata,and location of conodont and Protosalvinia samples. Filled circles give generallocation of samples listed in Appendix 1, open circles are reports from literaturementioned in the text. G-S-W-C � Genesee-Sonyea-Westfalls-Canadaway groups.B. General Famennian stratigraphy of the Appalachian and Illinois basins, Fa-mennian conodont zonation and relative duration from Kaufmann (2006), andrelative position of Protosalvinia Interval (circle pattern). Shaded strata are pre-dominantly black shale; T � position of Three Lick Bed. CZ � conodont zones:t � triangularis; c � crepida; r � rhomboidea; m � marginifera; t � trachytera;p � postera; e � expansa; p � praesulcata.

western New York and Pennsylvania, as well as in equivalentstrata of the Chagrin and Huron shales in Ohio, Protosalviniahave been reported from numerous sites (see Murphy, 1973). TheProtosalvinia-bearing interval is approximately 20 m thick.Above the strata containing Protosalvinia is the Lyons Road Bed,a phosphorite-rich shelly lag accumulation in the middle of theEllicott Shale that yielded a diverse conodont fauna; it includesBispathodus stabilis (Branson and Mehl, 1934a), Mehlina gradataYoungquist, 1945, Mehlina strigosa (Branson and Mehl, 1934a),Palmatolepis schindewolfi, Polygnathus homoirregularis Ziegler,1971, P. semicostatus Branson and Mehl, 1934a, Pelekysgnathusellicottensis new species, and P. inclinatus (localities E-G, Table1; Fig. 4). Hass (1958) reported Palmatolepis glabra distorta(specimens lost, Repetski, pers. comm.) and Polygnathus semi-costatus from the middle Ellicott Shale in Jamestown, NY (USGSSD 3948), and Polylophodonta confluens (Ulrich and Bassler,1926) in the Ellicott Shale just west of Jamestown (USGS SD3974). The upper range of Palmatolepis glabra distorta ends inthe Upper trachytera Zone and Polylophodonta confluens has a

similar range. The specimens assigned to Polygnathus homoir-regularis are similar to those illustrated by Sandberg and Ziegler(1979) from the Upper velifer Zone (�Upper trachytera Zone),and they gave a range of Upper velifer to Lower costatus zones(�Upper trachytera to Middle expansa zones). Based on cono-donts in Poland, Dzik (2006) suggested a slightly shorter range,trachytera Zone to Lower jugosus Zone (�Lower expansa Zone).Metzger (1989) reported P. homoirregularis in association withPalmatolepis postera and P. rugosa rugosa from the Sheffield-Maple Mill formations in eastern Nebraska, a faunal associationof the Lower postera through Middle expansa zones. Althoughconodonts of zonal significance have not been found with Pro-tosalvinia in the Ellicott Shale, the close occurrence of Palma-tolepis glabra distorta, Polylophodonta confluens, and Polygna-thus homoirregularis indicates the Upper trachytera Zone.

Protosalvinia in the Huron Shale in Ohio and Kentucky werefound in association with Palmatolepis glabra angusta and Bis-pathodus stabilis (localities H, I, Table 1). The KEP-#8 drill corecontains Protosalvinia at 380 feet, just above strata containingPalmatolepis glabra distorta, P. sigmoidea, P. rugosa trachyteraZiegler, 1960, and P. falcata (Figure 5). This association indicatesthe Lower-Upper trachytera zones. Palmatolepis perlobata max-ima Muller, 1956 and P. perlobata ssp. are found 40 m (130 feet)higher in the core within the upper black shale of the Three LickBed; P. p. maxima ranges from the Upper marginifera Zone intothe Lower expansa Zone, an indication that the Three Lick Bedis no higher than Lower expansa Zone.

In the Chattanooga Shale of the southern Appalachian Basin,Protosalvinia are associated with a disconformity that rests onstrata assigned to the Upper marginifera Zone, such as at Hurri-cane Bridge, a reference locality for the Chattanooga Shale (lo-cality L, Table 1; Over, 2007). Elsewhere, such as at ChestnutMound (locality K, Table 1), Protosalvinia are associated withthe conodonts Palmatolepis glabra distorta, likely reworked, andBispathodus aculeatus aculeatus, which marks the base of theMiddle expansa Zone, a higher occurrence, or suggests that Pro-tosalvinia remains may also be reworked.

In the Clegg Creek Member of the New Albany Shale Pal-matolepis glabra angusta occurs in great abundance above thefirst occurrence of Protosalvinia-bearing shale and a diverse co-nodont fauna characteristic of the Lower and Upper trachyterazones. Two drill cores from the Clegg Creek Member of the NewAlbany Shale in Indiana yielded Protosalvina and conodonts. Inthe 1–3 Kavanaugh core (locality D2; Table 1), the lowest oc-currence of Protosalvina is at or above a conodont-rich horizonthat contains Palmatolepis perlobata sigmoidea, P. perlobatagrossi, P. glabra distorta, and possibly Polylophodonta confluens.A conodont-rich horizon containing a similar fauna, likely at thesame disconformity, is found in outcrops in northern Kentucky(localities A and B, Table 1). The fauna here contain taxa that aretypical of the Upper marginifera Zone (e.g., Palmatolepis mar-ginifera utahensis, as well as those that range through this zone,like P. perlobata grossi, and P. rugosa rugosa; Figure 6). Theoccurrence of Palmatolepis glabra distorta and P. falcata abovethis widespread disconformity in the Walton 1–12 core (localityC2; Figure 7), as well as the even higher occurrence of P. glabraangusta, in which the entire element assemblage is preserved (lo-cality C3), indicates that the disconformity and Protosalvinia areno higher than the Upper trachytera Zone.

Four other reports have suggested the relative conodont-basedposition of the Protosalvinia Interval. Ettensohn et al. (1988)placed the Protosalvinia Interval in the Lower expansa Zonebased on one sample from the New Albany Shale in Kentucky(Table 1, locality B), using the conodont Polygnathus experplexusSandberg and Ziegler, 1979 as the key taxon, a conodont whichhas a reported range from the base of the Lower expansa Zone

72 JOURNAL OF PALEONTOLOGY, V. 83, NO. 1, 2009

FIGURE 2—Digital light microscope image of Protosalvinia Dawson showing typical occurrence, and scanning electron microscope (SEM) image that showsreticulate pattern. Scale bars are 1.0 mm. 1. OSU 52,998, shelly siltstone bed in medial Ellicott Shale, Chautauqua County, New York (locality H�), uncoated.2, OSU 52,999, Huron Shale, Delaware County, Ohio (locality I), coated.

TABLE 1— Conodont and plant taxa recovered from Protosalvinia-bearing and associated strata. See Appendix 1 for locality information. cc � Clegg CreekMember of New Albany Shale; es � Ellicott Shale Member of Chadokoin Formation; hs � Huron Shale Member of Ohio Shale; ug � Upper GassawayMember of Chattanooga Shale; Bi. � Bispathodus; Br. � Branmehla; M. � Mehlina; Pa. � Palmatolepis; Pe. Pelekysgnathus; Po. � Polygnathus.

Taxon/Location A B C1 C2 C3 D1 D2 D3 E F G H I J K Lunit: cc cc cc cc cc cc cc cc es es es es hs hs ug ug

Protosalvinia � � � � � � � � � �‘‘black spines’’ � � �Bi. aculeatus aculeatus � �Bi. stabilis � � � � � � �Br. inornata � � � � � � �M. gradata �M. strigosa � � �Pa. falcata � � � � � �Pa. glabra angusta � � � � �Pa. glabra distorta � � � � � � � �Pa. inflexa �Pa. inflexoidea �Pa. m. marginifera �Pa. marginifera utahensis �Pa. perlobata grossi � � � �Pa. perlobata maxima �Pa. perlobata sigmoidea � � � �Pa. rugosa rugosa �Pa. schindewolfi � � � � �Pandorinollina insita �Pe. inclinatus � � �Pe. ellicottensis � �Po. brevilaminus �Po. pennatulus � �Po. perplexus � � ?Po. homoirregularis �Po. planirostratus �Po. semicostatus � � � �Po. szulczewski �Polylophodonta confluens � � � � ? � �

A—K-I65–22, base of Protosalvinia, New Albany; B—NE-2 of Ettensohn lag below #10, New Albany; C1—Walton 1-12 Core, 4-1, 474.15 m, NewAlbany; C2—Walton 1-12 Core, 3-15, 472.78 m, New Albany; C3—Walton 1-12 Core, 2-14, 472.06 m, New Albany; D1—1-3 Kavanaugh Core, 8-9, 615.17m, New Albany; D2—1-3 Kavanaugh Core, 6-7, 613.75 m, New Albany; D3—1-3 Kavanaugh Core, 2-9, 610.94 m, New Albany; E—Hass (1958) SD3947,SD3948, SD3974, Ellicott Shale; F—Little Elk Creek, PA, Ellicott Shale; G—Lyons Road Bed in Chautauqua Creek, Chautauqua County, Ellicott Shale; H—medial Ellicott Shale, tributary to Clear Creek, Chautauqua County, Ellicott Shale; I—Huron Shale in S Delaware County, Ohio Shale; J—Huron Shale nearMorehead, KY, Ohio Shale; K—Chestnut Mound 6.3 m above base, Chattanooga Shale; L—Hurricane Bridge 9.75 m above base, Chattanooga Shale.


FIGURE 3—Ranges of selected conodonts found in association with Protosalvinia-bearing strata and zone defining taxa (darker lines). Ranges are fromKonigshof and Picha (1991), except: 4—Over (2007) and Oliver et al. (1969); 6—extension from Gutschick and Sandberg (1991); 8—Dreesen and Dusar(1974); 9a—Dreesen and Dusar (1974); 17—Sandberg and Ziegler (1979).

into the Middle expansa Zone. The sample also includes Pal-matolepis glabra distorta, P. falcata, P. perlobata grossi, Polyg-nathus brevilaminus Branson and Mehl, 1934a, Polygnathus pen-natulus Ulrich and Bassler, 1926, P. semicostatus, andPolylophodonta confluens, in which the occurrence of P. glaberadistorta and other taxa indicate a lower biostratigraphic positionor a reworked fauna. Considering the synonymy of P. experplexuswith P. perplexus (Metzger, 1989) the range of P. perplexus isfrom within the Lower trachytera Zone into the Upper expansaZone; Dreesen and Dusar (1974) indicated an even lower firstoccurrence of P. perplexus, in which specimens from Belgiumfirst occur in the Lower marginifera Zone. Thus, the report in-terpreted as within the Lower expansa Zone by Ettensohn et al.(1988) based on P. experplexus is questionable; other taxa suggestthe Upper trachytera Zone. The fauna at the disconformity as-sociated with the first occurrence of Protosalvinia from localityA suggests that the horizon is correlative with the disconformityat localities B, C1, and D2 (Table 1; Appendix 1).

The zonal position for Protosalvinia recovered 2 m above thebase of the Bakken Shale in northwest North Dakota from NDGSwell no. 4340, as reported by Holland et al. (1987), is problematicbecause the occurrence of conodonts from the lower Bakken inother cores relative to the occurrence of Protosalvinia is unclear.Lower Bakken conodonts described by Hayes (1985) and Hollandet al. (1987) from drill core include Bispathodus jugosus Branson

and Mehl, 1934a, Polygnathus experplexus, and P. granulosusBranson and Mehl, 1934a, that are interpreted as Lower expansaZone. Yet, the Trident Member of the Three Forks Formation inMontana, that is correlated to the Three Forks Formation thatunderlies the Bakken, contains a conodont fauna of the Uppervelifer Zone (�Upper trachytera Zone; Sandberg and Poole,1977; Holland et al., 1987). Thus the base of the Bakken Shaleand position of Protosalvinia are not well constrained. Discon-formities in the Upper Devonian black shales are often cryptic,and stratigraphic packets of similar lithologies can have irregulardistributions (e.g., Fuentes et al., 2002; Schieber and Over, 2005).The occurrence of Protosalvinia in the Lower Bakken is likelyno higher than the Lower expansa Zone, but it may be as low asthe Upper trachytera Zone.

Gutschick and Sandberg (1991) proposed a range for Proto-salvinia from the Upper rhomboidea Zone into at least the Lowerexpansa Zone and three separate blooms in the Michigan Basin.The lower age is based on correlation of gamma-ray logs withthe report of Protosalvinia by Matthews (1983) and was not sub-stantiated by a conodont fauna. In addition, Protosalvinia werereported in a core from the Michigan Basin of northern Indianafrom below a sample listed as Late or Latest marginifera (Sand-berg et al., 1994, table 2, sample NH90-3, p. 241), although bothconodonts listed, Palmatolepis marginifera marginifera and P.glabra distorta, range higher. These taxa also co-occur at locality


FIGURE 4—Digital SEM images of conodonts from Protosalvinia-bearing strata in Pennsylvania (locality F, 1–6, 8–14, 16–20), and New York (locality H,7, 15). Scale bar is 1.0 mm. 1, 2, Polygnathus homoirregularis Ziegler. 1, OSU 52,951, P1 upper view; 2, OSU 52,952, P1 upper view. 3, 4, Polygnathussemicostatus Branson and Mehl. 3, OSU 52,953, P1 upper view; 4, OSU 52,954, P1 upper view. 5, Palmatolepis schindewolfi Muller, OSU 52,955, P1 upperview. 6, 7, 12, 13, 16–20, Pelekysgnathus ellicottensis. 6, OSU 52,957, P1 inner lateral view; 7, OSU 52,959, P1 inner lateral view; 12, OSU 52,625, P1 lowerview; 13, OSU 52,958, P1 upper view; 16–18, OSU 52626–OSU 52,628, inner lateral views of conical elements; 19, OSU 52,630, P1 inner lateral view; 20,OSU 52,629, P1 lower view. 8, 10, 14, Mehlina strigosa (Branson and Mehl). 8, OSU 52,960, P1 inner lateral view; 10, OSU 52,961, P1 inner lateral view;14, OSU 52,962, P1 upper view. 9, 11, Mehlina gradata Youngquist. 9, OSU 52,963, P1 inner lateral view; 11, OSU 52,964, P1 inner lateral view. 15,Pelekysgnathus inclinatus Thomas, OSU 52,956, P1 inner lateral view.

A (Table 1) in association with an Upper trachytera Zone fauna.Palmatolepis gracilis sigmoidalis Ziegler, 1962b, which first ap-pears in the Upper trachytera Zone, was recovered 12 m higherin the core (Sanderg et al., 1994, p. 241). Conodonts from con-cretions in the Huron Shale at the type locality in northern Ohiothat included Palmatolepis gracilis sigmoidalis were used to es-tablish the occurrence of Protosalvinia in the Upper trachyteraZone. Also present were Bispathodus stabilis, Mehlina strigosa,P. glabra distorta, P. perlobata helmsi Ziegler, 1962b, P. perlo-bata sigmoidea Ziegler, 1962b, Polygnathus brevilaminus, andScaphignathus velifer Helms, 1959, similar to conodont faunas

from other Protosalvinia-bearing strata elsewhere in the Appala-chian and Illinois basins. Sandberg et al. (1994) went on to pro-pose that Protosalvinia from the Michigan Basin, and by impli-cation for other parts of the eastern United States, range from aslow as the rhomboidea Zone to the expansa zones.

In the Amazon Basin of northern Brazil, Protosalvinia are asso-ciated with a miospore flora of the upper VCo Zone that is equivalentto the ‘‘trachytera or more probably postera or even early expansaconodont Zones . . . ’’ (Loboziak et al., 1997). Protosalvinia foundwith Bispathodus aculeatus aculeatus Branson and Mehl, 1934a, atChestnut Mound in the upper Gassaway Member of the Chattanooga


FIGURE 5—Digital mirrored inverse SEM images of conodont molds inshale matrix, all upper views of P1-elements, Protosalvinia-bearing OhioShale strata from Kentucky Geological Survey core KEP-#8 drill core, corescale in feet; P � Protosalvinia. Scale bar is 1.0 mm, except for 3. 1, Pal-matolepis inflexoidea Ziegler, 418.8�, OSU 52,631; 2, Palmatolepis schinde-wolfi Muller, 407.5�, OSU 52,632; 3, Palmatolepis rugosa trachytera Ziegler,388.0�, OSU 52,633; 4, Palmatolepis glabra distorta Branson and Mehl,381.75�, OSU 52,634; 5, 7, Palmatolepis perlobata sigmoidea Ziegler. 5,385.3�, OSU 52635; 7, 383.5�, OSU 52,624. 6, Palmatolepis falcata (Helms),384.5�, OSU 52990. 8, Palmatolepis perlobata ssp., 249.3�, OSU 52,994. 9,Palmatolepis perlobata maxima Muller, 244.8�, OSU 52,995.

Shale indicate the Middle expansa Zone or higher, a reworked faunaland floral assemblage, or a higher occurrence as proposed by Gut-schick and Sandberg (1991). Protosalvinia have also been reportedfrom Montana in strata assigned to the Exshaw Shale (Cross, 1982in Matthews, 1983), which is considered equivalent, in part, to theThree Forks Formation and the Bakken Shale; the lower Exshaw atthe type locality is Lower-Middle expansa Zone (Richards and Hig-gins, 1989) although Macke (1993) has questioned the biostrati-graphic assignment of these strata.

CONCLUSION

Protosalvina in the Appalachian and Illinois basins have awidespread occurrence commonly associated with a horizon of

reworked conodonts in strata no higher than the Upper trachyteraZone. Reports of lower first occurrences in the rhomboidea ormarginifera zones are not well constrained by conodonts. Higheroccurrences in the Lower and Middle expansa zones are knownfrom the Chattanooga Shale in Tennessee in association with re-worked conodonts, and possibly from the Exshaw Shale. Proto-salvinia from the Bakken Shale in the Williston Basin are notwell constrained by conodonts, but have been associated withconodonts of the Lower expansa Zone.

SYSTEMATIC PALEONTOLOGY

The higher taxonomic categories of Sweet (1988) and elementnotation of Purnell et al. (2000) are followed herein. Figured spec-imens are housed in the Orton Museum at Ohio State University,Columbus, Ohio (OSU). All other materials are in the collectionsat SUNY-Geneseo stored by locality.

Class CONODONTI Branson, 1938Order PRIONIODONTIDA Dzik, 1976

Family ICRIODONTIDAE Muller and Muller, 1957Genus PELEKYSGNATHUS Thomas, 1949

PELEKYSGNATHUS ELLICOTTENSIS new speciesFigure 4.6, 4.7, 4.12, 4.13, 4.16–4.20

? Pelekysgnathus cf. P. brevis SANDBERG AND DREESEN, JOHNSTON AND

CHATTERTON, 2001, p. 21, pl. 4, figs. 17, 18 [only].

Diagnosis.⎯P1 element: segminate, very short blade, slightlyarched and curved, high inclined cusp, single broad, laterallycompressed, prominent vertical to anteriorly directed denticle onanterior end of blade; a small low denticle may be developedposterior of the large denticle. Basal cavity extends length of el-ement, two tips in expanded basal cavity beneath cusp. Coniformelements: nongeniculate, erect to proclined, asymmetrical to com-pressed base, ornamented with fine longitudinal striae.

Etymology.⎯Named for Ellicott Township, Chautauqua County, NewYork.

Material examined.⎯Locality F, H, 9 P1 elements, 5 conical elements.Types.⎯Holotype OSU 52959, the specimen illustrated in Figure 4.7 from

Little Elk Creek, PA, Locality F; paratypes OSU 52625–OSU 52630 fromsame location; OSU 52958 from tributary to Clear Creek, Locality H.

Discussion.⎯Insufficient coniform elements were recovered toreconstruct an apparatus. Large translucent specimens of Pelek-ysgnathus ellicottensis have numerous small denticle-like featuresdistinguished by white matter within the blade between the cuspand large anterior denticle; yet a small specimen (Fig. 4.6) doesnot have minor denticles developed. The P1 element of Pelekys-gnathus ellicottensis has a very short blade that is similar to Pe-lekysgnathus brevis Sandberg and Dreesen (1984), but the lateris characterized by two or three short denticles anterior of theprominent cusp. Johnston and Chatterton (2001) illustrated twosmall P1 elements bearing a prominent cusp and single or noposterior denticles similar to P. ellicottensis from lower Famen-nian strata of western Canada as P. cf. P. brevis; no coniformelements were described. Pelekysgnathus australis Nicoll andDruce 1979 from the high Famennian of Australia is characterizedby coniform elements that are typically striated; the P1 elementmay also have striae on the cusp, but has three to four denticles(Nicoll and Druce, 1979). ?Pelekysgnathus guizhouensis as re-constructed by Over (1992), a high Famennian taxon, has coni-form elements that have longitudinal striae; the P1 element differssignificantly in having numerous denticles on a longer blade.

ACKNOWLEDGEMENTS

Specimens used in this study were collected as part of NSF EAR-9614314to DJO and by FRE in studies of black shale strata NSF EAR-8513414.Thanks to the donors of the Petroleum Research Fund, administered by theAmerican Chemical Society (Grant #38523-AC8) for financial support; J.Rupp from the Indiana State Survey, P. Gooding and R. Daniel from theKentucky Geological Survey, and G. Schumacher from the Ohio Geological


FIGURE 7—Digital mirrored inverse SEM images of conodont molds inshale matrix, P1-elements from New Albany Shale Walton 1–12 drill core.Core boxes are numbered, conodont-bearing horizons are shown by horizontalbars and core interval of imaged conodonts is labeled; P � Protosalvinia.Scale bar � 1.0 mm, except figs. 8 and 9 where bar � 0.5 mm. 1, 6, Pal-matolepis glabra distorta Branson and Mehl. 1, 5–6, OSU 52,986, upperview; 6, 3–14, OSU 52,987, upper view. 2, Palmatolepis perlobata sigmoideaZiegler. 5–11, OSU 52,988, upper view. 3, Palmatolepis perlobata grossiZiegler, 5–11, OSU 52,989, upper view. 4, Polygnathus perplexus Thomas,5–11, OSU 52,991, upper view. 5, Polygnathus brevilaminus Branson andMehl, 5–11, OSU 52,992, lateral view. 7, Palmatolepis falcata (Helms),2–12, OSU 52,993, upper view. 8, 9 Palmatolepis glabra angusta Capkinoglu,2–14, OSU 52,996, OSU 52,997, upper views.

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FIGURE 6—Digital mirrored inverse SEM images of conodont molds in shale matrix, all upper views of P1-elements, from Protosalvinia-bearing strata atKI-65—22, 20.4 m above the base of the New Albany Shale (locality A). Scale bars are 1 mm—figures 5, 18–21 are larger scale. 1, Palmatolepis perlobatamaxima Muller, OSU 52,965. 2, 3, Palmatolepis glabra distorta Branson and Mehl, 2, OSU 52,966; 3, OSU 52,967. 4, Palmatolepis falcata (Helms), OSU52,968. 5, Palmatolepis glabra angusta Capkinoglu, OSU 52,969. 6, Polylophodonta confluens (Ulrich and Bassler), OSU 52,970. 7, Palmatolepis perlobatagrossi Ziegler, OSU 52,971. 8, Palmatolepis rugosa rugosa Branson and Mehl, OSU 52,972. 9, 10, 12, Palmatolepis perlobata sigmoidea Ziegler. 9, OSU52,973; 10, OSU 52,974; 12, OSU 52,975. 11, 14, 15, Palmatolepis schindewolfi Muller. 11, OSU 52,976; 14, OSU 52,977; 15, OSU 52,978. 13, Polygnathusplanirostratus Dreesen and Dusar, OSU 52,979. 16, Polygnathus szulczewskii Matyja, OSU 52,980. 17, Polygnathus pennatulus Ulrich and Bassler, OSU52,981. 18, 19, Palmatolepis marginifera marginifera (Helms). 18, OSU 52,982; 19, OSU 52,983. 20, 21, Palmatolepis marginifera utahensis Ziegler andSandberg, 20, OSU 52,984; 21, OSU 52,985.

Survey, for support of drill core sampling. C. Corradini, G. Klapper, and R.Metzger provided thoughtful and thorough reviews of the manuscript. Thisstudy is a contribution to IGCP 499—Devonian Land Sea Interaction: Evo-lution of Ecosystems and Climate.

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ACCEPTED 18 AUGUST 2008

APPENDIX 1

Locality information—same as localities in Table 1.

A—KI65-22, 20.4 m above base of New Albany Shale (BU-2 of Ettensohnet al., 1988)—roadside exposure at SW corner of junction of KY 245and I-65, 7.7 km south of Shepardsville, KY, Shepardsville Quad, UTM16S06152(E)41980(N), Bullet County, Kentucky.

B—NE-2 of Ettensohn et al. (1988), lag below #10 in road cut along east-bound lane of Bluegrass Parkway, 1.9 km east of the Exit 10 ramp ontoKY 52, Lebanon Junction Quad, UTM 16S 061892(E)417908(N), NelsonCounty, KY.

C—Walton 1–12 Drill Core, Indiana Geological Survey, Barr Township, Dav-iess County, Montgomery Quad, UTM 16S 0508095(E)4285090(N).

D—Kavanaugh 1–3 Drill Core, Indiana Geological Survey, Perry Township,Martin County, Loogootee Quad, UTM 16S 0494611(E)4276705(N).

E—Hass (1958): SD3947—type Ellicott Shale, creek exposure west of James-town, NY, just north of Hunt Road, 1.2 km west of intersection withSouthwestern Drive, Lakewood Quad, UTM 17T 06422(E)46610(N),Chautauqua County, New York; SD3948—road side exposure 2.5 mabove curb at southeast corner of Willard and Pardee streets in James-town, NY, Lakewood Quad, UTM 17T 06475(E)46619(N), ChautauquaCounty, New York, SD3974—float from railroad exposure on Conrail(Erie) Railroad between Hunt and Bigtree roads 1 km southwest of Lak-ewood, NY, Lakewood Quad, UTM 17T 06365(E)46610(N), ChautauquaCounty, New York.

F—Little Elk Creek, medial Ellicott Shale, 10 cm-thick shell bed in creek exposurebelow Franklin Center Road bridge, west of Franklin Center, PA, AlbionQuad, UTM 17T 05595(E)46437(N), Erie County, Pennsylvania.

G—Lyons Road Bed in Ellicott Shale, stream exposure at Lyons Road bridgeover Chautauqua Creek, 2.7 km northwest of Summerdale from junctionwith NY 430, Sherman Quad, UTM 17T 06159(E)46756(N), ChautauquaCounty, New York.


H—medial Ellicott Shale, stream exposure in tributary to Clear Creek, S ofHarris Hollow Road at approximately 1780�, Gerry Quad, UTM 17T06517(E)46762(N), Chautauqua County, New York.

H�—shelly siltstone bed with Protosalvinia, stream exposure in Folsom Creekat approximately 1790�, N of Gerry Ellington Road, 5.5 km NE of Gerry,New York, Gerry Quad, UTM 17T 06487(E)46755(N), ChautauquaCounty, New York.

I—High Banks Metro Park, north of Columbus, Ohio, east off US 23—creekexposure north of nature center, Powell Quad, UTM 17T03273(E)44464(N), Delaware County, Ohio.

J—I-64 near Morehead, KY (RW-1 of Ettensohn et al. (1988)—7 m abovebase of Huron Member of Ohio Shale—roadside exposure on I-64 east-bound, 0.3 km east of KY 1722 (Lower Licking Road) bridge, FarmersQuad, UTM 17S 02728(E)42272(N), Rowan County, Kentucky.

K—Chestnut Mound 6.3 m above base of Chattanooga Shale—roadside ex-posure on US 70N, 1.6 km west of junction with TN 53 in ChestnutMound, Buffalo Valley Quad, UTM 16S 06043(E)40076(N), SmithCounty, Tennessee.

L—Hurricane Bridge (DE-2 of Ettensohn et al., 1988), 9.75 m above base ofChattanooga Shale—roadside exposure north of Smithville, TN 56, 2.5km west of Hurricane Bridge, Center Hill Dam Quad, UTM 16S06124(E)39899(N), DeKalb County, Tennessee (reference section forChattanooga Shale).

KEP-#8 Drill Core, Kentucky Geological Survey, CC 5800� FSL � 1600�FEL Sec. 3-Y-75, 4 km (2.5 miles) SE of Vanceburg, KY on KY 546(AA) 4.2 km (2.6 miles) east of intersection with KY 59, VanceburgQuad, UTM 17S 03006(E)42729(N), Lewis County, Kentucky.

protosalvinia dawson and associated ......b. general famennian stratigraphy of the appalachian and...

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