Late Devonian Goniatites (Cephalopoda, Ammonoidea) from New

Bulletins of American PaleontologyNumber 374, July 2008

Late Devonian Goniatites (Cephalopoda, Ammonoidea)

from New York State

M. R. House and W. T. Kirchgasser

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Bulletins of American PaleontologyNumber 374, July 2008

Late Devonian Goniatites (Cephalopoda, Ammonoidea)

from New York State

M. R. House and W. T. Kirchgasser

ISSN 007-5779ISBN 978-0-87710-476-6

Library of Congress Catalog Card Number 2008931960© 2008, Paleontological Research Institution, 1259 Trumansburg Road, Ithaca, New York 14850, U. S. A.

Michael House (at far right) on a “Friends of Devonian” fi eld trip, western New York, 1966, with (left to right) John Huddle (U. S. Geological Survey), Ed Buehler (SUNY Buff alo), Larry Rickard (N. Y. State Geological Survey), Bill Kirchgasser (SUNY Potsdam), and Jon Harrington (Cornell University). Photograph by Prof. J. W. Wells (Cornell University).

Michael House (second from left) with Tony Dunn (SUNY Potsdam) and Prof. and Mrs. J. Wells, at Luck Stone Lodge, Sheldrake Point, Cayuga Lake, 1984.

Michael House (seated at right) with Gil Klapper (at left, University of Iowa) and Th omas Becker (standing, Southampton University and Universität Münster), in Western Australia, 1990.

TRIBUTE

MICHAEL ROBERT HOUSE

(1930-2002) Michael House was raised on the Mesozoic ammonites along the coast of his native Dorset. While studying at Cambridge, he found Devonian goniatites on the rocky shores of Cornwall and Devon and began a lifelong pursuit of their relatives around the globe, collecting bed by bed and tracking down type specimens in museums large and small, making friends everywhere. He came to America to fi nd Professor John Wells at Cornell University in Ithaca, New York, and was guided to the Naples area where the seed that grew into this monograph was planted.

House was a tireless ambassador for the Devonian with an ever-expanding goniatite portfolio, a true internationalist, a Friend of the Devonian and all its students, profes-sional and amateur. He was a dedicated teacher and mentor to generations of students at the universities at Durham, Oxford, Hull and Southampton.

Among the many awards and tributes is a volume of papers in his honor (Becker & Kirchgasser, 2007) that includes a list of his one-hundred-plus publications.

R. T., & W. T. Kirchgasser, eds. 2007. Devonian Events and Correlations. Geological Society (London) Special Publication 278, 280 pp.

LATE DEVONIAN GONIATITES (Cephalopoda, Ammonoidea)FROM NEW YORK STATE

M. R. House* &W. T. Kirchgasser

Department of Geology, State University of New York at Potsdam, Potsdam, New York13676, U. S. A., email [email protected] [*deceased]

ABSTRACTThis paper is a revision of the classic nineteenth century goniatite collections from the Devonian of New York State de-scribed by James Hall and John M. Clarke. Type material is illustrated photographically, mostly for the first time; thetaxonomy is revised, and ontogenetic and statistical data are given, where possible. Detailed biostratigraphic work is re-ported through the late Givetian, Frasnian, and Famennian, aimed at determining the precise horizon and location ofprimary material at 184 localities. A description of the considerable additional goniatite material is integrated with thatof the classic type specimens to treat the faunas systematically and as a whole. Noteworthy is the different contributionto knowledge provided by a range of preservation patterns, especially of the material preserved as barite replacements fromthe Cashaqua Shale. The goniatite-bearing horizons are mostly associated with transgressive pulses that punctuate theCatskill Delta succession. The major ones are the upper Tully Limestone, the Geneseo and Renwick black shales, the Ge-nundewa Limestone, and the Middlesex, Rhinestreet, Pipe Creek, and Dunkirk black shales.

The refined goniatite succession reported is perhaps the most detailed known in the world through the interval stud-ied. The New York faunal levels are correlated with those known elsewhere, especially with Old World successions, andwith the new international standard established by Becker & House (2000). Minor regional variations give a moredetailed Appalachian regional zonation of 25 zones, which is integrated with the international standard. The successionassociated with the major late Givetian faunal turnover (Taghanic Event) is described, and the influence of sea levelchanges and anoxia on faunal entry and diversification is reviewed. Very little endemicity is recognized in the faunas, apartfrom the multilobedTriainoceratidae that are well developed in the mid-Frasnian of central New York. The late Frasniansuccession associated with the Lower and Upper Kellwasser Events is documented. Parallel conodont work provides cor-relation with the conodont zonation scale. New taxa described are Koenenites styliophylus kilfoylei n. ssp., Koenenites beckerin. sp., Sandbergeroceras? enfieldense n. sp., and Manticoceras sinuosum clausium n. ssp.; the older names sinuosum Hall,1843, and tardum Clarke, 1898, are regarded as subspecies ofManticoceras sinuosum (Hall, 1843). Lectotypes are desig-nated for Pharciceras tridens (G. & F. Sandberger, 1850), Sandbergeroceras syngonum Clarke, 1897, Acanthoclymenia ge-nundewa (Clarke, 1898), Chutoceras nundaium (Hall, 1874), Koenenites styliophilus styliophilus (Clarke, 1898), K.?fasciculatus (Clarke, 1898),Manticoceras contractum Clarke, 1898,M. sinuosum apprimatum Clarke, 1898,M. sinuosumtardum Clarke, 1898,M. accelerans Clarke, 1897, Carinoceras sororium (Clarke, 1898),Delphiceras cataphractum (Clarke,1898), Sphaeromanticoceras rhynchostomum (Clarke, 1898), S. oxy (Clarke, 1897), Tornoceras uniangulare uniangulare(Conrad, 1842), and Truyolsoceras bicostatum (Hall, 1843). The Geneseo Limestone Horizon is proposed as a new in-formally named horizon of the Genesee Group.

1HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES

INTRODUCTIONNowhere in the world is the Upper Devonian exposed socontinuously and accessibly as in New York State. The lowerUpper Devonian (upper Givetian-Frasnian) in particular isseen in quite unparalleled excellence, and this interval hasthe greatest potential for refining the biostratigraphic recordof goniatite cephalopods. The Upper Devonian in New YorkState crops out over an area of approximately 15,000 mi2

(39,000 km2). The sequences are well exposed in innumer-able creek, gully, and riverside sections. The beds are undis-

turbed, and over a wide outcrop tract more than 280 mi(450 km) long, the dip of the rocks rarely exceeds 1.5o tothe south. With so extensive an outcrop, and so simple astructure, the facies changes from the thin marine sequencenear Lake Erie on the west to the thick non-marine se-quences in the Catskill Mountains to the east can be corre-lated and analyzed in great detail (Text-fig. 1).

These goniatite faunas have been known since the workof James Hall (1843) and Lardner Vanuxem (1839, 1842),but it was their description by John M. Clarke (1898,1899a, b) that made them particularly well known. Clarke

discovered very well-preserved material, including rich bar-itized faunas that enabled extraction by etching, and he de-veloped techniques for producing artificial molds of materialfrom siltstones. His descriptions of the early stages of manyforms, particularly species of Manticoceras, are still amongthe more interesting of goniatite ontogenetic studies. Thisearly work was largely collated and revised by A. K. Miller(1938) in a review of American Devonian ammonoids, buthardly any of Clarke’s material was reillustrated photograph-ically, and the original drawings were merely reproduced.Modern illustration has long been needed.

The New York goniatite sequence has largely been ig-nored biostratigraphically, apart from broad reviews byHouse (1962, 1978). The early faunas described by Clarkewere from isolated localities, the relations of which have notbeen understood, and in addition he used a stratigraphic ter-minology that has long been out-of-date. Indeed, the posi-tion of many of the localities was given in so cursory amanner as to make their geographical position doubtful ex-cept in the most general terms.

This study was commenced by the authors in 1964, pri-marily to relocate the old sources of material and to establishtheir stratigraphic relations. This was an ambitious under-taking and involved the study and measuring of many sec-tions in western upstate New York to document the facieschanges and discover the sequence of ammonoid faunas.

The framework was built on the lithological correlations es-tablished by G.W. Colton, J. F. Pepper, W. deWitt, Jr., andR. G. Sutton in a series of contributions. So excellent arethe exposures, however, that individual goniatite horizonshave been traced, in some cases, for distances of more than50 mi (80 km). Substantial detail has thus been added tothe stratigraphical picture built up by many earlier geolo-gists.

This work describes the occurrence and sequence of go-niatite faunas for the lower Upper Devonian which is un-equalled in the world for its stratigraphical discrimination.It contributes to a detailed documentation of goniatite evo-lution during this period and adds detail on the facieschanges associated with the goniatite horizons as they passeastward toward the Catskill Delta.

HISTORICAL REVIEWWhen, in the autumn of 1836, James Hall and LardnerVanuxem commenced field work in the Devonian area forthe newly created Geological Survey of New York, much hadalready been accomplished in the establishment of thestratigraphy and distribution of the Devonian rocks, eventhough the System was not itself designated until 1839(Sedgwick &Murchison, 1839a: 259; 1839b: 121). The pi-oneer period of geological investigation, up to the establish-

Text-fig. 1. Map of upstate New York showing the outcrop area of late Devonian rocks (Tully Limestone and younger) (stippled), the 15'quadrangle map areas, and (inset) the location of the diagram. The Finger Lakes are marked in black. In addition to the state quadranglemaps, current editions of the New York State Atlas and Gazeteer (Anonymous, 1993) are a convenient source of maps for site location.

2 BULLETINS OF AMERICAN PALEONTOLOGY no. 374


ment of the Survey, has been reviewed in a thorough essayby J. W. Wells (1963).

With the publication of Part III of the Geology of NewYork (Vanuxem, 1842), and particularly of Part IV (Hall,1843) with its geological map and systematic stratigraphicaltreatment, a firm foundation of geological knowledge wasestablished. Understandably, at that time, correlation withEurope was imprecise, and Hall thought that the Portageand Chemung rocks, now referred to the Upper Devonian,preceded the Old Red Sandstone. Their Devonian age seemsfirst to have been appreciated by Ferdinand Roemer in 1846(Clarke, 1921: 172; Wells, 1963: 64) who had visited Hallat Albany in 1845.

Hall had, however, recognized the broad pattern ofthickening from the marine rocks of the west to the terres-trial facies in the east, and had recognized the extent of theTully Limestone, and of the major wedges of black shales, allgiving time horizons extending eastward to the thick Catskillred-bed succession. Hall described Devonian goniatites froma horizon within the Tully Limestone up to faunas on LakeErie shore in Chautauqua County, now known to belong toequivalents of the German Nehdener Schiefer (early Fa-mennian). The source of this material will be discussed inthe next section.

Hall had the benefit of accompanying Charles Lyell overmuch of the ground covered by his survey before publicationof The Geology of New York, Part IV (Friedman, 1998). Lyellwas the leading theoretical geologist of his time and we haveno way of assessing the contributions he made to the devel-opment of Hall’s ideas and vice versa. It is a matter of surprisethat Hall did not consider with any accuracy the matter ofcorrelation with Europe. By the time Hall met him, Lyellhad already, as President of the Geological Society of Lon-don, presided over many of the bitter arguments related tothe establishment of the Devonian System in Europe (Rud-wick, 1985). There is no evidence in Hall’s 1843 volumethat these problems had been resolved, nor of their relevancefor the North American rocks. Only a correlation with theOld Red Sandstone was attempted, and that was relatedsolely to the terrestrial Catskill facies.

Vanuxem (1842), in reporting on the third geologicaldistrict to the east of Hall’s fourth district, was already inrocks of a more clastic facies, but he too described goniatitesfrom rocks that would now be termed Frasnian, or lowerUpper Devonian. New names were given to other goniatitesin the same year by Conrad (1842), one of the state geolo-gists whom Hall replaced as State Paleontologist in 1843.

So far as goniatites are concerned, investigations ceasedfor a substantial time, apart from spasmodic records (Hall,1874, 1875). During the course of his Paleontology of NewYork, Hall (1876, 1879; Hall & Clarke, 1888) described ad-

ditional goniatites, but with the arrival of J. M. Clarke atthe Survey, substantial advances followed (Clarke 1882,1883, 1884, 1885). In 1885, Clarke described some mate-rial from the Cashaqua Shale and discussed the goniatites(Clarke 1891a, b). The supposed recognition of a clymeniid(Clarke, 1892) was mistaken, but the baritic preservation ofthe material might well have led him to appreciate the valueof acid-etching of New York material. Certainly it was thedevelopment of these techniques that resulted in his impor-tant study of goniatites in The Naples Fauna (Clarke 1898,1899a, b). In this work much new material was described,especially from the Cashaqua, Angola, and Hanover Shales,but the locality details were poorly recorded; a useful sourceof information on localities is a small guidebook by Clarketo fossiliferous localities in western New York (Clarke,1899c). Had stratigraphical precision been given for the go-niatite localities and faunas described by Clarke, there is lit-tle doubt that he would have laid the foundation of Frasniangoniatite biostratigraphy. As he did not do this, that distinc-tion went to Denckmann, Frech, and especially Wedekind,working on the much less well-exposed rocks in the RhenishSchiefergebirge of Germany.

The early decades of the twentieth century saw manynew records of goniatites, mostly in the bulletins accompa-nying the geological quadrangle mapping of upstate NewYork. New forms, however, were not recognized. Loomis(1903) described early stages of goniatites from the LeicesterPyrite. Clarke (1902, 1903) added further notes, and thesecond part of The Naples Fauna was published in 1904.The next significant discovery was of Paralegoceras? milleri(nowMaeneceras milleri) by Flower & Caster (1935) in theConewango Group in neighboring Pennsylvania, althoughit was not until a later paper (Miller & Flower, 1936), thatthe generic assignment to Sporadoceras was given and itsrecognition as an indicator of the mid-Famennian was ap-preciated.

A useful compilation and revision of earlier work of NewYork Devonian goniatites was given by A. K. Miller in 1938.This took advantage of the substantial German work of theprevious 30 years to give more up-to-date generic assign-ments, but no fieldwork was done for this compilation, andmost illustrations were merely copied from earlier sources,especially Clarke’s monographs of 1898 and 1899.

Subsequently, Koenenites was recognized in the WestRiver Shale (Wells, 1956b), Cheiloceras in the GowandaShale, and Pharciceras in the Tully Limestone (House,1962). The latter two genera are key markers in the Euro-pean biostratigraphy. Clarke’s supposed clymeniid wasshown to be a manticoceratid (House, 1961) and a broadcorrelation of North American and European goniatitezones was attempted (House, 1962, 1968, 1978). The

tornoceratid succession of New York formed the basis for anallometric study of the group (House, 1965). Substantialnew information on the biostratigraphic occurrences of Pro-beloceras and Ponticeras was given in a review of these generaby Kirchgasser (1968, 1975), and details have been given ofgoniatite levels in the Genesee Group (Kirchgasser, 1973,1982, 1985, 1994, 1996a, b, 1998) and Sonyea Group(Cashaqua Shale) (Kirchgasser, 1965, 1967, 1969, 1975)and comparisons made with the conodont succession(Kirchgasser, 1974, 1994; Klapper et al., 1995). Finally, thepresent authors published general reviews of Frasnian faunasof New York (Kirchgasser & House 1981; House & Kirch-gasser, 1993) that contributed to the establishment of a newinternational goniatite zonation for the Frasnian Stage(Becker et al., 1993, Becker & House, 2000).

STRATIGRAPHYINTRODUCTION

This section reviews the levels at which late Devonian am-monoids have been discovered in New York State. To locatethe exact horizons of earlier collections, considerable effortwas expended in measuring sections across the western part

of the State. This led to some revisions of correlations. Nev-ertheless, the stratigraphic framework is built on the detailedwork of G. W. Colton, W. de Witt, Jr., J. F. Pepper, and R.G. Sutton and the reviews of L. V. Rickard. The relation ofour sequences to earlier work is discussed in the followingtext. Our locality numbers are shown in Text-fig. 4 andlisted inTable 1. Horizon numbers and letters follow the lo-cality numbers as in “Loc. 34/6a.” Localities in an alphabet-ical series (as in Loc. R, I-2) refer to earlier sectiondescriptions, notably by de Witt and Colton. For localityreferences, we used the U. S. G. S. quadrangle maps; a usefulmodern guide to the localities is the New York State Atlasand Gazeteer (1993 and later editions).

The terminology for the main lithostratigraphic units inwestern and central parts of New York State is shown inText-fig, 2, which is based on the review of Rickard (1975).See also the reviews of Oliver et al. (1968, 1969) and Oliver& Klapper (1981a).

We have mostly been successful in locating the exact lev-els of the famous faunas described by J. M. Clarke (1898,1899a, b) with the notable exception of the strange fauna ofearly beloceratids, here placed in the genusNaplesites, whichwas recorded from Naples (Ontario County) in a facies thatappears to be within the Rhinestreet Shale (see Addendum).

A comment is required on measurements. The currenttopographic maps show only scale in miles and altitude infeet. Thus we are obliged to give locations using Englishmeasurements, but metric equivalents are given in paren-theses. For measured sections, metric measurements aregiven, with English equivalents in parenthesis. For paleon-tological descriptions only metric measurements are given.

A facies transect for the rocks with which we are con-cerned is shown in Text-fig. 3. An outline from this (Text-fig. 4) is used to indicate the most important of the sectionswe have studied; those of earlier workers are referred to inthe text, as appropriate. Fuller details of localities, includingrelevant 7.5’ quadrangle maps and county location, are givenin Table 1.

TULLY FORMATION

The Tully Limestone is one of the main regional, base-linemarker units for the interval examined here. TheTully is thelast of the substantial calcareous intercalations that charac-terize marine Devonian in New York State. The name wasapplied by Vanuxem (1839: 278; 1842: 285) for the se-quence near Tully (Onondaga County) and nearby areas.Near Tully it is over 9 m (30 ft)-thick (Trainer, 1932). East-ward there is a passage into more clastic facies (Cooper &Williams, 1935; Cooper, 1968; Johnson & Friedman, 1969)and westward, although anomalously thick owing to a “reef”or mound complex at Borodino, the thicknesses are reduced

Text-fig. 2. Lithostratigraphic divisions of late Devonian rocks ofNew York State (based on Rickard, 1975), with chronostratigraphicboundaries of the stage divisions drawn following decisions of the In-ternational Commission on Stratigraphy (ICS) and InternationalUnion of Geological Sciences (IUGS).


Text-fig.3.Faciesdiagramindicating

themainrockunitsinthelateDevonianofNew

YorkStatefrom

which

goniatitesaredescribedinthisstudy.Attopleft,readCorell’s(for

Correll’s)PointG

oniatiteBed.


Text-fig.4.O

utlineofthemainrockunitsinthelateDevonianofNew

YorkStateandtheposition

ofsomeoftheprincipallocalitiesandsectionsusingthenumbering

system

followedinthetext.


Table 1. Locality numbers used in this study showing the quadran-gle maps and counties for the locality areas, The widely availableNew York State Atlas and Gazetteer (1:150,000; DeLorme Map-ping) is a useful modern guide to the topo maps and localitieswithin the state. An asterisk indicates that the section is illustratedin one of the text-figures. Localities with letter abbreviations of15’ quads and numbers are U. S. G. S. localities from de Witt &Colton (1978).

No. Locality 15’ Quad 7.5’ Quad CountyTULLY FORMATION1 Beards Creek Caledonia Leicester Livingston2 Gorham Phelps Rushville Ontario3 Lodi Glen Ovid Lodi Seneca4 Grove’s Creek Qry Genoa Sheldrake Seneca5* Carpenter Falls Skaneateles Spafford Cayuga6 Borodino Skaneateles Spafford Onondaga7* June’s Quarry Tully Tully Onondaga8* Tinkers Falls Tully Tully Cortland9a New Lisbon Hartwick Morris Otsego9b Laurens Hartwick Mt. Vision Otsego9c Lebanon Morrisville West Eaton Madison

GENESEE GROUP10* Lake Erie Silver Creek Angola Erie11* Eighteenmile Crk Eden Eden Erie12* Cazenovia Creek Depew Orchard Park Erie13* Cayuga Creek Depew East Aurora Erie14* Elevenmile Creek Attica Corfu Genesee14a Murder Creek Attica Alexander Genesee15* Linden Batavia Batavia South Genesee15a* Linden Falls Batavia Batavia South Genesee16* White Creek Batavia Stafford Genesee17* Taunton Gully Caledonia Leicester Livingston17a Spezzano Caledonia Leicester Livingston18* Beards Creek Caledonia Leicester Livingston18a* Beards Creek Caledonia Leicester Livingston19* Fall Brook- Caledonia Geneseo Livingston

Dewey Hill19a* Fall Brook Caledonia Geneseo Livingston20* Hemlock Honeoye Honeoye Livingston

(Big Tree Rd)20a Hemlock Honeoye Honeoye Ontario

(Huff-Barnard Rd)21 Abbey Gulf Honeoye- Honeoye- Ontario

Canandaigua Bristol Center22 Barnes Gully Canandaigua Canandaigua- Ontario

Lake23* Seneca Point Creek Naples Middlesex Ontario23a Seneca Point Creek Naples- Middlesex- Ontario

Canandaigua Canandaigua-Lake23b Genundewa Point Naples Middlesex Yates24 Bethany Center Batavia Batavia So. Genesee

(embankment)24b Bethany Center Batavia Batavia So. Genesee

(roadcut)25 Fir Tree Point Watkins Reading Ctr Yates

(Wg-2-3)25a Baskin Pt (Ov-4) Ovid Dundee Yates26a* Whetstone Brook Honeoye Honeoye Ontario27 Lodi Glen Ovid Lodi Seneca27a Romulus Town Ovid Ovid Seneca

Quarry

Table 1 (continued).No. Locality 15’ Quad 7.5’ Quad County27b Hubbard Quarry Genoa Sheldrake Seneca28 Pleasant Valley New Berlin Sherburne Chenango29a Hicks Point Naples Middlesex Ontario46c Snyder’s Gully Naples Middlesex Yates-

OntarioA-23b Genundewa Point Naples Middlesex Yates

(Nap-4)B Shuman Cemetary Phelps Rushville Yates

(Ph-5)C Sartwell Ravine Penn Yan Penn Yan Yates

(Py-11)D Lakeside Country Penn Yan Penn Yan Yates

Club (Py-11a)E Willow Grove Penn Yan Keuka Park Yates

Crk (Py-14)F Sunset Point Penn Yan Keuka Park Yates

(Py-16)F1 0.9 mi South Penn Yan Keuka Park Yates

CrosbyG Gully N of Ovid Dundee Yates

Plum Pt (Ov-3a)H-25a N of Baskin Pt Ovid Dundee Yates

(Ov-4)I S of Baskin Pt Ovid Dundee Yates

(Ov-5)J N side of Fir Tree Watkins Glen Reading Ctr Yates

Pt (Wg-2)K Green Pt (Wg-7a) Watkins Glen Reading Ctr SchuylerL Mill Creek (Ov-10) Ovid Dundee-Lodi SenecaM N of Tommy Ovid Ovid Seneca

Creek (Ov-13)N-27a Romulus Town Ovid Ovid Seneca

Quarry (Ov-16)O-27b Hubbard Quarry Genoa Sheldrake Seneca

(Gen-2)P Ovid-Shedrake Genoa Sheldrake Seneca

Road (Gen-1a)Q Glenwood Crk (I-1) Ithaca Ithaca West TompkinsR Williams Brook Ithaca Ithaca West Tompkins

(I-2)S S of Williams Ithaca Ithaca West Tompkins

Brook (I-2a)T Gulf Creek (I-11d) Genoa Ludlowville TompkinsU Esty Glen (I-11-b) Ithaca Ithaca West TompkinsV Twin Glens at Ithaca Ithaca West Tompkins

McKinneys (I-11a)W S of McKinneys Ithaca Ithaca West Tompkins

(I-11c)X Fall Creek, Dryden Ithaca East Tompkins

Ithaca (Dy-10)Y Baker Quarry Dryden Ithaca East Tompkins

(Dy-10a)Z Cascadilla Creek Dryden Ithaca East Tompkins

(Dy-10b)AA South Hill, Ithaca Ithaca West Tompkins

Ithaca (Dy-10c)BB Enfield Glen Ithaca Ithaca West Tompkins

(I-8a)CC Big Salmon Crk, Genoa Genoa Cayuga

Genoa (Gen-13a)DD Big Salmon Crk Genoa Genoa Cayuga

Venice Ctr (Gen-14a)


Table 1 (continued).No. Locality 15’ Quad 7.5’ Quad CountyDD1 Cornwall Gully Penn Yan Penn Yan Yates

(Py-10)EE Willow Grove Penn Yan Keuka Park Yates

(PY-14)FF Sunset Point Penn Yan Keuka Park Yates

Gully (Py16)GG Creek E of Ovid Dundee Schuyler

Starkey (Ov-6)HH Granger Point Naples Middlesex Ontario

(Nap-3)II(XX) 0.8 mi S of Middle- Naples Middlesex Yates

sex (Nap-8)JJ Chidsey Point Penn Yan Pulteney Steuben

Creek (Py-7)KK-F Sunset Point Penn Yan Keuka Park Yates

Gully (Py-16)LL-L Mill Creek (Ov-10) Ovid Lodi SenecaMM Sheldrake Creek Ovid Ovid Seneca

(Gen-1a)NN Coy Glen (I-3) Ithaca Ithaca West TompkinsOO Lick Brook (I-8) Ithaca Ithaca West Tompkins

SONYEA GROUP30* Eighteenmile Crk Eden Eden Erie30a Pike Creek Silver Crk & Angola & Erie

Eden Eden31 Smoke Creek Buffalo Buffalo SE Erie32* Cazenovia Crk Depew Orchard Park Erie33* Cayuga Creek Attica Cowlesville Erie &

Wyoming34* Murder Creek Attica Alexander Genesee35* Little Tona- Batavia Batavia South Genesee

wanda Creek37 Wyoming Gulf Batavia Wyoming Wyoming38* Beards Creek Caledonia Leicester Livingston39* Buck Run Creek Nunda Sonyea & Livingston

Mt Morris40* Cottonwood Wayland Conesus Livingston

Point Gully41 Shurtleff ’s Gully Honeoye Livonia Livingston41a North McMillan Wayland Conesus Livingston

Creek42* Whetstone Brk Honeoye Honeoye Ontario43* Briggs Gully Wayland & Springwater & Ontario

Naples Bristol Sprs44* Randall Gully Canandaigua Bristol Center Ontario44a Barnes Gully Canandaigua Canandaigua Ontario

Lake45 Hicks Point Naples Middlesex Ontario46 Cook Point Naples Middlesex Ontario46a Long Point Naples Middlesex Ontario46b Whiskey Point Naples Middlesex Ontario46c Snyder’s Gully Naples Middlesex Yates &

Ontario46d Griesa Hill Road Naples Middlesex Ontario46e Bristol Springs Rd Naples Bristol Sprs Ontario46f Bristol Springs Naples Bristol Sprs Ontario

Road Gully47* Conklin Gully Naples Middlesex Yates47a Rumpus Hill Naples Middlesex Yates47b Clark Gully Naples Middlesex Yates

Table 1 (continued).No. Locality 15’ Quad 7.5’ Quad County47c Lee Cemetery Naples Middlesex Yates47d School No. 7, Naples Middlesex Yates

Middlesex47e Lower East Hill Rd Naples Middlesex Yates47f School No. 2, Naples Middlesex Yates

Italy Valley48* Wagener Glen Penn Yan Pulteney Steuben49 Bobell Hill Oxford Brisben Chenango

WEST FALLS GROUP50 Sturgeon Point Silver Creek Angola Erie51 Big Sister Creek Silver Creek Angola Erie52 Eighteenmile Creek Eden Eden Erie53 Hampton Brook Eden Hamburg Erie54* Cazenovia Creek Springville Colden Erie55* Hunter Creek Depew East Aurora Erie56 Sheldon Creek Arcade Strykersville Wyoming57* Varysburg Attica Attica Wyoming58* Johnson Creek Attica Attica Wyoming59a* Tannery Brook Attica Attica Wyoming60* Relyea Creek Portage Warsaw Wyoming61* Stony Creek Portage Warsaw Wyoming63* Kennedy Gulf Batavia Dale Wyoming64 Harford Mills Harford Harford Tioga65a* Parrish Gully Naples Middlesex Yates65* Buck Run Creek Nunda Mount Morris Livingston66 Hungerford Quarry Dryden Ithaca East Tompkins67 Fairfield Forest Dryden Speedsville Tioga68 Bald Mtn Dryden Speedsville Tompkins69 Owego Owego Owego Tioga70 Elmira Elmira Elmira Chemung71* Lake Erie Shore Silver Creek Angola Erie

(Farnham Creek)72* Point Breeze Silver Creek Angola Erie73* Big Sister Creek Silver Creek Angola Erie

(Angola)74* Hampton Brook Eden Hamburg Erie75* Cazenovia Creek Springville Colden Erie77* Glade Creek Arcade Strykersville Wyoming78* Sheldon Creek Arcade Strykersville Erie &

Wyoming79* Varysburg Attica Attica Wyoming80 Varysburg Attica Attica Wyoming

(Gassman Road)81* Johnson Creek Attica Attica Wyoming82* Relyea Creek Portage Warsaw Wyoming83* Stony Creek Portage Warsaw Wyoming84* Kennedy Gulf Batavia Dale Wyoming85 Wolf Creek Portage Portageville Wyoming86* Genesee Gorge Portage Portageville Wyoming &

Livingston87 Stony Brook Glen Wayland Dansville Steuben87a I-390–Dansville Wayland Dansville Steuben88 Eden Valley Eden Hamburg Erie89a* Walnut Creek Silver Creek Silver Creek Chautauqua89* Walnut Creek Silver Creek Silver Creek Chautauqua90* Irish Gulf Eden & Hamburg & Erie

Springville Colden91* Glade Creek Arcade Strykersville Wyoming92* Beaver Meadow Arcade Strykersville Wyoming

Creek (Java)


until it is last seen on the eastern side of Canandaigua Lakeat Gage Gully. Farther westward, its place is taken by theLeicester Pyrite, a diachronous remanié deposit marking thecoalescence of horizons at the compound unconformity be-tween the Hamilton and Genesee Groups: pre-Tullydisconformity (Taghanic Unconformity), intra-Tully discon-

formity (Lower-Upper Tully boundary), and post-Tully dis-conformity (Taghanic Onlap of black Geneseo Shale overgray Moscow Member of the Windom Shale) (Brett &Baird, 1982, 1994, 1996, 1997; Baird & Brett, 1986a, b,2001). The detailed lithostratigraphy of the Tully has beenthoroughly described by Heckel in his excellent paper

Table 1 (continued).No. Locality 15’ Quad 7.5’ Quad County93 Wiscoy Creek Portage Portageville Allegany94 Bald Hill, Dryden Speedsville Tompkins

Caroline95 N Chemung Waverly Waverly & Chemung

Wellsburg96 Van Etten Waverly Van Etten Chemung

CANADAWAY GROUP97* Corell’s Point Dunkirk Brocton Chautau-

qua98* Little Cana- Dunkirk Brocton Chautau-

daway Creek qua99* Walnut Creek Cherry Creek Forestville Chautau-

qua100 Smith Mills Cherry Creek Perrysburg & Chautau-

Forestville qua101 Big Indian Creek Silver Creek Farnham Cattarau-

gus102 Little Indian Creek Silver Creek Farnham Cattarau-

gus

Table 1 (continued).No. Locality 15’ Quad 7.5’ Quad County103 Cattaraugus Creek Cattaraugus Gowanda Reserva-

(tributary on tionCat. Indian Res.)

104 Cattaraugus Creek Cattaraugus Gowanda Cattarau-(S Branch) gus

105 Clear Creek Eden North Collins Erie(North Branch),Taylor Hollow

106 Clear Creek Eden N Collins & Erie(North Branch), LangfordMarshfield

107 Anthony Gulf Springville Colden Erie108 Gears Creek Springville Holland Erie109 Java Arcade Strykersville Wyoming

CONNEAUT AND CONEWANGO GROUPS110 Porter Creek, Clymer Sherman Chautau-

Summerdale qua111 Howard Cambridge

Quarries Springs Edinboro N Erie (PA)

Text-fig. 5. Diagram showing the units within the Tully Limestone recognized by Heckel (1973).


(1973) (Text-fig. 5), and summarized by Heckel (1997).Building on Heckel’s work, Baird & Brett (2003) and Bairdet al. (2003) have added significant detail to the correlationsand faunal relations within the Tully, particularly in theSherburne-Oneonta area of Otsego and Chenango counties.

Goniatites are known from a single horizon in theMoravia Bed of the Upper Tully and from black shale inter-calations in the Filmore Glen Bed of the uppermost Tully.The horizon in the Moravia Bed is in the upper part of BedG of West Brook Member of Cooper & Williams (1935:791) at Tinkers Falls and June’s Quarry, near Tully (House,1968: 1065) and at corresponding levels farther west (Text-fig. 6). The type locality for Pharciceras amplexum (Hall,1886: pl. 127 (12), fig. 1) was given as Lodi Landing, SenecaLake. J. W.Wells located a probable in situ representative ofthis form 0.38 m (1.25 ft) above the Bellona Coral Bed, inLodi Glen, at the same horizon within the Upper Tully as atTully and Tinkers Falls. Apart from P. amplexum, tornocer-atids occur at these localities and some have been referred toTornoceras cf. arcuatum House (1965: 108).

Indeterminate crushed molds of probable pharciceratidsare known from the black shale interbeds of the FillmoreGlen Member of the Upper Tully between Skaneateles andCayuga lakes. Westward from Cayuga Lake the Upper Tullyunits are progressively removed by erosional beveling (Text-figs. 5, 7), and locally the Leicester Pyrite, with tornocer-atids, is found at the Tully-Geneseo contact and, west ofCanandaigua Lake, at the Moscow-Geneseo contact. At

Gorham (Ontario County) between Seneca and Canan-daigua lakes (Loc. 2), the apparent easternmost occurrenceof the Leicester Pyrite, indeterminate tornoceratid nucleioccur in pyritic nodules at the sharp contact between theBellona Coral Bed of the Tully and the black Geneseo Shale(House, 1965: 83). West of Canandaigua Lake, where theTully is missing, the tornoceratids in the Leicester Pyrite atthe base of the black Geneseo Shale that are referable toTornoceras uniangulare uniangulare (Conrad, 1842) wereprobably all exhumed as pyrite debris from the underlyingMoscow Shale (Baird & Brett, 1986a, b).Localities.–Locality numbers with asterisks have sections il-lustrated in Text-fig. 6. The one goniatite horizon foundwithin the Tully Limestone proper is the level yielding Phar-ciceras amplexum. Also listed here are some localities of theLeicester Pyrite (Loc. 1) or localities where goniatites areknown close to theTully (Locs 2 and 4) or, in the east, wheregoniatites apparently occur in equivalents of the Tully (Loc.9). Detailed correlation was given by Heckel (1973, 1997)for units of the Tully (Text-fig. 5); see also Baird & Brett(2003) and Baird et al. (2003).

Locality 1, Beards Creek.–The Leicester Pyrite occurs atthe base of the section illustrated in Text-fig. 8 (Loc. 18) atBeards Creek, Leicester (Livingston County), 0.1 mi (0.16km) west of the railroad crossing, at an altitude of 638 ft(194.5 m). The Leicester Pyrite (Text-fig. 8) here forms dis-continuous lenses of pyrite nodules and steinkerns. Thepyrite is also exposed in several neighboring creeks. This is

Text-fig. 6. Diagram showing sections with Pharciceras bed in the Tully Limestone. For other localities see text.



the source area for the types of Tornoceras uniangulare uni-angulare.

Locality 2, Gorham.–Along Flint Creek, at Gorham(Ontario County), there are exposures of the Tully Lime-stone, which here is 1.5 m (5 ft) thick. Exposures were wellseen in 1959 east of East Swamp Road, south of Gorham,during excavations for draining the swamp where a pyriticlevel at the top yielded Tornoceras (House, 1965: 83).

Locality 3, Lodi Glen.–On the eastern side of SenecaLake, at Lodi Point Landing (Seneca County), is the localitygiven by Hall (1886: pl. 127 (12), fig. 1) as the source ofthe type material of Pharciceras amplexum. In Lodi Glen,above the Landing, J. W. Wells located in situ a probablePharciceras, 0.38 m (1.25 ft) above the Bellona Coral Bed(Heckel, 1973: 40) in the Tully Limestone. The relationshere are as at Tinkers Falls (Loc. 8).

Locality 4, Grove’s Creek Quarry.–Southeast of Ovid(Seneca County) at Grove’s Creek Quarry, pyritic tornocer-atids referred to Tornoceras aff. uniangulare occur below theTully Limestone (House, 1965: 83) in the upper part of theunderlying Moscow Shale. This is within the Gorge GullySubmember of the Windom Shale (Brett & Baird, 1994).

Locality 5*, Carpenter Falls.–At Carpenter Falls, belowthe bridge on Appletree Point Road over Bear Swamp Creek,1 mi (1.6 km) north of New Hope (Cayuga County), thelevel of the Bellona Coral Bed is 0.48 m (1.6 ft) below thebridge foundation and the top of the unit above is the ex-pected level for Pharciceras (Text-fig. 6, Loc. 5).

Locality 6, Borodino.–Tornoceras is recorded from theWest Brook Member of the Tully (House, 1965: 83) atBorodino (Onondaga County; Heckel, 1973: 45). We havenot located the horizon in situ. It is probable that this is fromthe same level as other Tully goniatite faunas.

Locality 7*, June’s Quarry.–One mi (1.6 km) northeastof Tully (Onondaga County), June’s Quarry (or Ravine) hasprovided by far the richest collection from the Pharciceraslevel (Text-fig. 6, Loc. 7). Determinable material was firstfound here by W. A. Oliver, Jr. The best collecting is fromthe old quarry surface on the western side, in the middlepart of the Moravia Bed of Heckel (1973: 41), 1.27 m (4.2ft) above the base of Bed G of the West Brook Member ofCooper & Williams (1935). It should be noted that thePharciceras level is not in the Apulia Member of Cooper &Williams (1935), as originally thought (House, 1962: 274).

Locality 8*, Tinkers Falls.–Tinkers or Tinker Falls (Cort-land County), 5 mi (8 km) east of Tully, is the type sectionfor the divisions of Cooper & Williams (1935: 791). Herethe Pharciceras bed was located within the Moravia Bed ofHeckel, approximately 1.47 m (4.8 ft) above the base of theWest Brook Member, in Bed G of Cooper & Williams(Text-fig. 6, Loc. 8), at the same level as in June’s Quarry.

Locality 9a, New Lisbon.–Goniatites occur in more clas-tic equivalents of theTully Limestone near New Lisbon (Ot-sego County). In the northern fork of Stony Brook, 1.5 mi(2.4 km) northeast of New Lisbon, crushed goniatites (poortornoceratids) occur in equivalents of theWest BrookMem-ber of Cooper & Williams (1935) and presumably in thevinicity of the projected level of theWest Brook in the NewLisbon section illustrated by Heckel (1973: fig. 19). GordonBaird (pers. comm., 2002) reported that the West BrookShale is deeply covered in the northern tributary but is stillwell exposed in the south branch. This is locality 34 of Bairdet al. (2003).

Locality 9b, Laurens.–At approximately 7 mi (11.3 km)southeast of New Lisbon (Otsego County), Pharciceras am-plexum occurs (USNM 96545) in the West Brook Memberand this is the most easterly record of the genus in the state.This locality is 1 mi (1.6 km) northeast of Laurens (OtsegoCounty). This is locality 37 of Baird et al. (2003).

Locality 9c, Lebanon.–There is a single specimen ofPharciceras amplexum recorded from theWest Brook Mem-ber (USNM 96552) in a ravine 0.5 mi (0.8 km) southwestof Lebanon (Madison County).

Other Localities.–In this and subsequent entries localitiesrefer to those given in previous publications and follow theprevious abbreviations.

Locality Gen-1.–From the black shales of the FillmoreGlen Bed of the uppermost Tully, at Sheldrake Creek, aboveCayuga Lake (western side), Gordon Baird and CarltonBrett collected several molds with well-preserved growthlines of probable Pharciceras, one of which preserves ventro-lateral furrows and a flattened venter but it does not show asuture. DeWitt & Colton (1978: locality Gen-1, table 1, pl.3) recorded the base of the Tully at 604 ft (184.1 m) eleva-tion, which is above Blew Road west of NY Rte. 89, approx-imately 1.5 mi (2.4 km) southwest of Sheldrake (SenecaCounty).

There are collections at the Paleontological Research In-stitution (PRI) of similar specimens of probable Pharcicerasfrom the Fillmore Glen Beds at Mack Creek (Loc. Gen-1c),near Yarnell Road, 2.6 mi (4 km) northeast of Ovid (SenecaCounty; locality 421 of J.W.Wells), at Fernbank (Loc. Gen-5a), along the western shore of Cayuga Lake, 0.5 mi (0.8km) north of Taughannock Creek (Tompkins County; local-ity 106c of J. W. Wells, Cornell University Museum of Pa-leontology [CU] G-89-92); section illustrated by Heckel(1973: pl. 2), de Witt & Colton (1978: pl. 3) and Grasso etal. (1986: fig. 8), and Brett & Ver Straeten (1997: 250–251,fig. 35).

GENESEE GROUP

This group shows a spectacular thickening eastwardfrom 2.7 m (9 ft) at Lake Erie, nearly 40 m (128 ft) in theGenesee Valley, 274 m (900 ft) at Cayuga Lake (de Witt &Colton, 1978), and perhaps as much as 490 m (1,600 ft) inthe Gilboa-Kaaterskill area (Rickard, 1975). In the west, be-tween Lake Erie and Canandaigua Lake, the Genesee Grouprests directly on the Moscow Shale of the Hamilton Group(Text-figs 3, 7). At the disconformable junction of the twogroups, the Leicester Pyrite, containing lenses of nodule-and steinkern-bearing detrital pyrite, is observed. It is a bednamed for exposures in the Genesee Valley at Leicester (Liv-ingston County). Huddle (1974, 1981) has demonstratedfrom conodonts that the Leicester Pyrite becomes progres-sively younger in a westward direction. There is strong evi-dence that the Leicester is a lag, or remanié, unit thatincludes exhumed material from the Moscow Shale (Baird& Brett, 1986a, b). The rich fauna of Tornoceras uniangulareuniangulare from the Leicester Pyrite (House, 1965) cannotunequivocally be assigned to the Moscow Shale, but it is dis-similar from tornoceratid faunas known in theTully Forma-

tion and lower Geneseo Shale, and the earlier assignment isthought to be the more probable. Eastward fromCanandaigua Lake, the Genesee Group disconformablyoverlies units of the Tully Formation (Heckel, 1973: pl. 2).

As presently interpreted, the Genesee Group comprisesbeds between the base of the black Geneseo Shale and thebase of the black Middlesex Shale (Text-fig. 3). The historyof the use of these terms is complex. Vanuxem (1842: 168)first used the term Genesee Slate, and his definition atCayuga Lake indicates that he restricted it to what is nowcalled the Geneseo Shale (a term which dates from Chad-wick, 1920). Vanuxem indicated that the Genesee Slate wasthe “Black Shale and Slate, and Upper Black Slate of Re-ports,” indicating how uncertain was the use of these terms.The present-day practice extends the use of the Genesee ei-ther in the form of a Formation (de Witt & Colton, 1959,1978) or as a Group (Chadwick, 1933; Rickard, 1975), thelimits embracing the sequence from the Geneseo Shalethrough the West River Shale, inclusive. This usage essen-tially dates from Clarke & Luther (1904). The various unitsand key marker-beds within have been discussed in detail

Text-fig. 7. Facies relations of upper Moscow Shale to lower West River Shale. Alignment of key horizons and the ammonoid and conodontsequences in the upper Moscow and Tully Formations and lower and middle Genesee Group, in central and western New York. Principalremanié or lag deposits include pyrite beds (in bold) and bone calcarenites (North Evans Limestone). Modified from Huddle (1981), Brett& Baird (1985), and Kirchgasser et al. (1994) and here plotted against the conodont and goniatite zonations. Conodont zonation afterKlapper (1989), Klapper & Johnson (1990), Ziegler & Sandberg (1990), and Klapper & Becker (1999). The change in age of the LeicesterPyrite westward was first demonstrated by Huddle (1974, 1981).



by Brett & Baird (1982, 1986, 1994, 1997), Baird & Brett(1986a, b), de Witt & Colton (1959, 1978), Baird et al.(1989), House & Kirchgasser (1993), Kirchgasser (1975,1985, 1994), Kirchgasser & House (1981), Kirchgasser etal. (1986, 1994), Kirchgasser, Brett, & Baird (1997), andKirchgasser, Over & Woodrow (1994, 1997). In a series ofpapers (references above) modeled on their work on the un-derlying Hamilton Group (Brett & Baird, 1996), CarltonBrett and Gordon Baird described the role of submarine ero-sion and sedimentary hiatuses associated with eustatic andtectonic cyclicity in the formation of the black shale, pyrite,and carbonate marker-beds of the Genesee Group. Thesehorizons are the keys to correlations within the eastwardthickening and coarsening facies of the group. Many ofthem are goniatite-bearing beds or proximal to goniatite-bearing beds, which makes them crucial to determining thegoniatite sequence within the group (see details in the lo-cality descriptions below).

In westernmost sections, west of Cazenovia Creek (Text-fig. 8, Loc. 12), the Leicester Pyrite is missing and the NorthEvans Limestone (Rickard, 1964) or “Conodont Bed” ofHinde (1879), a 75-100 mm (3-4 in) calc-arenitic bone-bedoccupies the interval between the upper Moscow Shale ofthe Hamilton Group and the overlying upper GenundewaLimestone of the middle Genesee Group or Formationalong the great Taghanic Unconformity (Text-fig. 7). Tracesof the horizon are found beneath the upper GenundewaLimestone as far east as the Genesee Valley (Kirchgasser,1996a, 1998; Kirchgasser & Koslowski, 1996; Kirchgasser& Vargo, 1998).

The North Evans is a winnowed remanié deposit (Baird& Brett, 1986a, b) famous for its abundant fish debris (Hus-sakof & Bryant, 1918; Turner, 1998). Moreover, it is thefamed “conodont bed” of Hinde (1897). Whereas the unitat its type area at Eighteenmile Creek (Loc. 11) (Text-fig.8) appears to occupy the time-rock interval between theupper Givetian varcus Zone and lower Frasnian MN (Mon-tagne Noire) Zone 2 (Text-fig. 7), recent taphonomic work(including that of Kirchgasser, 2001, 2004) suggests thatmost of the microvertebrate and conodont elements in theNorth Evans, throughout its geographic extent, are of lowerFrasnian (MN Zone 2) age, including the youngest zonalindicator in the unit, Ancyrodella recta Kralick, 1994; MNZone 3 begins in the overlying upper Genundewa Lime-stone with the entry of an early form of A. rugosa Branson& Mehl, 1934. Recently, pyritic protoconchs and earlywhorls of probable Frasnian goniatites (?Koenenites) havebeen recovered from the North Evans, which supports theview that the unit mostly records a sub-upper Genundewadisconformity with parts of the upper Penn Yan Shale andnodular lower Genundewa reworked or removed (Kirch-

gasser, 2002).

Geneseo Shale, Penn Yan Shale and Associated UnitsIn the type area of the Genesee Valley, the Geneseo Shale(Member) consists of two major tongues of black shale sep-arated by a sequence of dark gray shales (Text-fig. 8, Locs17-19; de Witt & Colton, 1978: pls 2, 6). The lower, orbasal tongue, is a continuous sequence of black shale, 1.2-1.5 m (4-5 ft) thick and overlain by 5.2-6.7 m (17-22 ft) ofdark gray shale.

The succeeding upper tongue consists of a 1.2-2.1 m (4-7 ft)-thick interval with a pair of black shales, separated bygray shale, which defines the top of the Geneseo Shale. Theupper tongue thins westward to a single black shale horizonat Cayuga Creek (Text-fig. 8, Loc. 13). Thus defined, theGeneseo Shale thickens from 1.1 m (3.5 ft) at Cayuga Creekto approximately 9 m (30 ft) in the Genesee Valley (Text-fig.8, Locs 17-19). In approximately the middle of the grayshale interval between the lower and upper tongues of theGeneseo Shale in the Genesee Valley, there is a prominent0.18-0.20 m (7-8 in)-thick argillaceous limestone contain-ing crushed Ponticeras. This level, earlier mistaken for theLodi Limestone (Kirchgasser, 1975), is informally namedherein the Geneseo Limestone Horizon. It can be tracedwestward, where the upper part of the gray shale interval ap-pears to have been progressively eroded, to Cayuga Creekwhere its position is in the gray shale beneath the uppertongue of the Geneseo Shale.

East of the Genesee Valley, the position of the GeneseoLimestone Horizon is difficult to recognize as the gray shaleinterval pinches out and the lower and upper tongues of theGeneseo Shale cannot be differentiated in the eastwardthickening succession of mostly continuous black shale withscattered concretionary (septarian) horizons (Text-fig. 3; deWitt & Colton, 1978: pls 2-3, 5). At Canandaigua Lake,the Geneseo Shale is 13 m (44 ft) thick at Menteth Gully (deWitt & Colton, 1978). Farther east, at Seneca and Cayugalakes, Baird & Brett (1986b) and Baird et al. (1989) recog-nized the Fir Tree Limestone (and equivalent Fir Tree Pyrite)as a submember in the upper Geneseo Shale. The Fir TreeLimestone occurs at the base of the Hubbard Quarry Shale,a submember of dark gray shale, comprising the uppermostdivision of the Geneseo Shale (Text-fig. 7). At the spectac-ular Taughannock Falls on the western side of Cayuga Lake,the Geneseo Shale is nearly 60 m (183 ft) thick and its toplies just below the lip of the falls (Baird & Brett, 1986b: 60-61, fig. 12).

Goniatites are poorly known in the Geneseo Shale andmost specimens are crushed flat with the shells having beenremoved in the dysoxic to anoxic basinal environments ofdeposition. These include rare Pharciceras? sp. at localities

Text-fig. 8. Sections of the lower Genesee Group comprising the Geneseo Shale, Penn Yan Shale, and Genundewa Limestone between Lake Erie andHemlock Lake, with the locality numbers used in the text. For Loc. 19/6 (LH) read barite (for baryte).



in Chenango and Otsego counties. Tornoceras sp. occursabundantly at 1.4 m (4.5 ft) (Text-fig. 8, Bed 18/1) and 5.1m (16.75 ft) (Bed 18/2) above the Leicester Pyrite at BeardsCreek (Loc. 18) in the Genesee Valley, but only one speci-men is known with a sharply projecting ventrolateral furrowthat could suggest a pharciceratid (from Bed 18/1). The oc-currence of Ponticeras in the Geneseo Limestone Horizon ofthe Genesee Valley has been noted above. Epitornoceras en-ters in the succession, as E. cf. mithracoides (Frech, 1887),approximately 12.2 m (40 ft) below the top of the GeneseoShale at Lodi Glen, Seneca Lake (Loc. 27) (House, 1965:119), 24-27 m (80-90 ft) above the top of the Tully Forma-tion. A specimen of E. cf. peracutum (Hall, 1879) comesfrom near the top of the Geneseo Member at HubbardQuarry (Loc. 27b) (House 1965: 119) on the western sideof the Cayuga Lake valley. The holotypes of E. peracutumand E. aff. peracutum (see House, 1978: 60) are from theIthaca Formation. Pyritic fragments and protoconchs of go-niatites (Tornoceras?, Ponticeras?) have been noted by us inthe residual pyrite of the Fir Tree Limestone in the SalmonCreek Valley east of Cayuga Lake (Locs CC, Gen-13a; DD,14a). Ponticeras perlatum (Hall, 1874) enters the successionin the dark gray to black shales at the top of the GeneseoShale (Hubbard Quarry Shale), the best locality being Hub-bard Quarry (Loc. 27b), which has yielded many specimens,although most are badly crushed; rarely are they in verticalorientation (Pl. 4, Fig. 5).

Penn Yan Shale and its Eastern EquivalentsThe Lodi Limestone (Lincoln, 1895), exposed in Lodi Glenon Seneca Lake (Loc. 27), is a 0.23 m (9 in)-thick concre-tionary bed 1.2 m (4 ft) above the top of the Geneseo Shale.The Lodi Limestone lies in the lower part of the local PennYan Shale (or Sherburne Siltstone). At this locality, at theRomulus Town Quarry, Ovid (Loc. 27a), and at HubbardQuarry (Loc. 27b) above Cayuga Lake, the Lodi bed hasyielded fine specimens of Ponticeras perlatum, the type ofwhich came fromHomer (Cortland County), farther to theeast (House, 1965; Kirchgasser, 1975).

The main bed of the Lodi Limestone at Lodi Glen is oneof several recurring horizons of impure carbonates with aulo-porid corals and other benthic fossils in the eastward thick-ening wedge of Penn Yan or Sherburne facies in the Seneca,Cayuga, and Skaneateles valleys (Baird & Brett, 1986a, b;Baird et al., 1989). On the western side of Seneca Lake theserecurring Lodi beds, designated Lodi Beds A-D, and similarhorizons higher up, can be seen in the bluff exposures (Locs25, 25a, Ov-5, Wg-2,Wg-7a); Lodi Bed B is the main Lodibed and Lodi Bed A is at the Geneseo/Penn Yan-Sherburnecontact. Some of the horizons have yielded Ponticeras butthey have not been systematically searched. On the eastern

side of Seneca Lake, a Lodi horizon (Lodi Bed C), 5.5 m(18 ft) above the main Lodi Bed B, yielded the large speci-men of P. cf. perlatum figured herein (Text-fig. 26D-E, Loc.M, Ov-13). The Lodi and post-Lodi horizons are the prob-able source of the records of P. perlatum, many in beds withaulaporid corals (Cladochonus beds or zones), in the Sher-burne Siltstone localities of Kindle (1896) and Boekenkamp(1963) at the south end of Cayuga Lake (Locs I-1, I-11a–d).

Westward (basinward) from Seneca Lake, the Lodi bedsconverge to a condensed sequence of reworked pyrite-phos-phate lags beneath black shales. In the Honeoye Lake Valley,on the western side of the regional basin axis, Lodi Bed Breappears as a single auloporid-bearing carbonate horizoncapped by a black shale designated Penn Yan Black Shale A,a horizon which marks the conodont Middle/Upper Devon-ian boundary in the region (Baird & Brett, 1986b; Baird etal., 1989; Kirchgasser et al., 1989; Kirchgasser, 1994). A sec-ond black shale (Penn Yan Black Shale B), approximately 2m (6 ft) above the Lodi at Abbey Gulf (Loc. 21) in the Ho-neoye Valley, has yielded Ponticeras.

Farther to the west, in the Genesee Valley, the LodiLimestone is represented by a single bed of nodules, withPonticeras perlatum, in the lower Penn Yan Shale approxi-mately 1 m (3 ft) above the top of the Geneseo Shale (Text-fig. 8, Loc. 18). Immediately above the Lodi is a pair of thinblack shales that represent Penn Yan Black Shales A and B.These post-Lodi black shales converge westward to a singleblack shale unit, thus constraining the top of the Lodi as farwest as Cayuga Creek where the Lodi horizon is representedby a thin, grey mudstone (Text-fig. 8, Loc. 13).

A thin black shale layer (Schumacher Bed [SB]), overly-ing a concretion band 4-5 m (12-17 ft) above the Lodi hori-zon in the Genesee Valley, is a key horizon for positioninggoniatite occurrences higher in the Penn Yan Shale andequivalent Sherburne Siltstone and the Ithaca Shale andSandstone. The SB black shale (Kirchgasser et al., 1986:249) and underlying concretion bed have been traced west-ward to Cayuga Creek (Text-fig. 8, Loc. 13). It is the posi-tion of the SB black shale at Honeoye and Canandaigualakes that constrains the position, in western sections, of theRenwick Shale, a major black shale marking the top of theSherburne Siltstone and base of the Ithaca Shale and Sand-stone in the Seneca and Cayuga valleys. At Abbey Gulf (Loc.21) in the Honeoye Lake Valley, the western tongue of theSherburne Siltstone, the Renwick Shale, and the informalunpublished “Abbey Gulf Beds” of G. Baird and C. Brett(Baird et al., 2006: 355; Zambito et al., 2007: 94, fig. 4),with indeterminate goniatites, form a sequence in the nearly9 m (30 ft) interval between the Lodi Limestone (Lodi BedB) and the SB black shale. Thus, the many records of Pon-ticeras perlatum reported from the top of the Sherburne and

BULLETINS OF AMERICAN PALEONTOLOGY no. 37416

base of the overlying Renwick Shale at localities at the southend of Cayuga Lake are stratigraphically below the SB blackshale. These are the highest confirmed records of P. perla-tum.

Higher in the Cayuga Lake Genesee Formation succes-sion, in the lower third of the Ithaca Shale and Sandstone,Chutoceras nundaium (Hall, 1874) occurs in the QuarrySandstone interval at Fall Creek, Ithaca (Loc. Dy-10), 72 m(235 ft) above the base of the Renwick Shale. It also occursin the same interval at South Hill, Ithaca (Loc. Dy-10c)(Kirchgasser, 1985), and possibly also in theWilliams BrookCoquinite atWilliams Brook (Loc. I-2) on the southwesternside of Cayuga Lake. Unfortunately, none of these horizonshas been found to the west of Cayuga Lake, although C.nundaium is represented in the equivalents of the upperPenn Yan Shale at Mill Creek, Seneca Lake (Loc. L, Ov-10).

Koenenites enters the succession in the thin styliolinidlimestones that characterize the upper Penn Yan Shale. Themost productive horizon is the Linden Horizon (Kirchgasser& House, 1981) in which Koenenites styliophilus styliophilus(Clarke, 1898) is locally common and can be well preserved.This horizon has been traced from Cayuga Creek (Loc. 13)in the west to the section above the Quarry Sandstone inter-val at Fall Creek (Loc. Dy-10), Ithaca, at Cayuga Lake(Kirchgasser, 1985).

Koenenites styliophilus styliophilus is also locally commonand well preserved in the Crosby Sandstone of Torrey et al.(1932) in sections in the vicinity of Keuka Lake (Locs C,Py-11, D, Py-11a, F, Py-16). At Keuka Lake, the CrosbySandstone marks the contact between the Penn Yan Shaleand the overlying Ithaca Shale and Sandstone. To the westat Seneca Point Creek, Canandaigua Lake (Loc. 23), Koe-nenites occurs in the “hiatus concretion horizon” of Baird(1976), a bed with the distinctive conodont fauna of theCrosby, which is found 7 m (23 ft) below the base of theGenundewa Limestone. To the east at Mill Creek, SenecaLake (Loc. L, Ov-10), the Crosby lies 6.1 m (20 ft) abovethe Linden Horizon and above other higher horizons alsoyielding Koenenites. Although the Crosby Sandstone appearsto occupy the stratigraphic position of the GenundewaLimestone east of Canandaigua Lake in some respects (seede Witt & Colton, 1978: pls 3-5), we believe it is an olderhorizon in the interval between the Linden Horizon and theGenundewa. The upper Penn Yan Shale appears to be trun-cated in the region of Keuka Lake and it could be that therethe Crosby Sandstone cuts down to the level of the LindenHorizon and incorporates Linden goniatites as a basal de-posit. No goniatites or fossils other than plant fragments areknown from the Crosby farther east. At Fall Creek, CayugaLake (Loc. Dy-10), the prominent siltstone immediatelyabove the Linden Horizon (Bluestone Bed of Kirchgasser,

1985) could conceivably be the Crosby Horizon. In the LickBrook section (Loc. I-8) of deWitt & Colton (1978: pl. 3),the Bluestone-Crosby bed would be the siltstone unit im-mediately below the covered interval in approximately themiddle of the Ithaca.

Acanthoclymenia enters the succession in the upper PennYan in Bed 15/9 on top of the Linden Horizon at Linden(Genesee County; Loc. 15) and there is also a record in theuppermost Penn Yan Shale at Sunset Point, Keuka Lake(Loc. F, Py-16), 0.30 m below the Crosby Sandstone.

Localities.–Spectacular eastward thickening is shown by thepre-Genundewa Genesee Group. On the accompanying di-agrams (Text-fig. 8), therefore, different scales are used forthe columns. The overlying Genundewa Limestone, whichis thin in relation to the lower levels, is also shown at an in-creased scale for clarity; the Genundewa is informally sub-divided into a nodular-bedded lower Genundewa and awell-bedded upper Genundewa. The nodular-bedded lowerGenundewa is missing in western sections between Linden(Loc. 15) and Lake Erie (Loc. 10). Locality numbers with as-terisks have sections illustrated in Text-figs 8-9.

Locality 10*, Lake Erie.–On the shore of Lake Erie, thereis a complete section of the Genesee Group, from the upper-most Windom Member of the Moscow Shale (HamiltonGroup) to the base of the Middlesex Black Shale (SonyeaGroup). This section is seen in the cliffs 170 yd (155 m)south of the mouth of Pike Creek, 1.4 mi (2.25 km) north-east of Jerusalem Corners (Erie County; Text-fig. 8, Loc.10). No goniatites have been collected there. A 31 mm (1.25in) unit at the base consists of laterally connected calcareniticlenses, 6 mm (0.25 in) thick [North Evans Limestone withmm-thick lenses of pyrite (?Leicester Pyrite) at its base;Kirchgasser, 2004], a black shale, 19 mm (0.75 in) thick,and a Styliolina-rich bed, 6 mm (0.25 in) thick at the top.Above this unit is 0.23 m (9 in) of interbedded very darkgray and black shale that represents undifferentiated PennYan Shale. The upper Genundewa Limestone lies above. Anearby section at the mouth of Pike Creek was illustratedby de Witt & Colton (1978: pl. 2).

Locality 11*, Eighteenmile Creek.–Along EighteenmileCreek, 137 yd (125 m) downstream from the westernmostof two railroad bridges, on the northern side of the creek,1.2 mi (1.9 km) northwest of North Evans (Erie County),there is an even more condensed sequence (Text-fig. 8, Loc.11). Here, beneath the upper Genundewa Limestone, is a se-quence of 6 mm (0.25 in) of black shale beneath 86 mm(3.4 in) of North Evans Limestone, followed by 25 mm (1in) of very dark gray to black shale representing the PennYan Shale. Pyritic specimens of ?Koenenites have recentlybeen noted in microfossil residues from the 29-mm-thick

HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 17

Bed 11/1e at the top of the North Evans Limestone (Kirch-gasser, 2002). No other goniatites are known from this lo-cality, which is a classic section of Grabau [1898–1899,Section I (H); illustrated by deWitt & Colton (1978: pl. 2)and Kirchgasser (1981: 11, fig. 1, reillustrated by Brett etal., 1997: 178, fig. 7)].

Locality 12*, Cazenovia Creek.–In Cazenovia Creek, asimilar section is seen, but in different facies. One mile (1.6km) south of Spring Brook (Erie County), and below theintersection of North Davis and Conley roads, a westwardflowing tributary joins Cazenovia Creek, and the sectiongiven here (Text-fig. 8, Loc. 12) is 20 yd (18 m) downstreamof the confluence. A thin lens (6 mm or 0.25 in) of LeicesterPyrite is followed there by 0.66 m (2.2 ft) of black and darkgray shale (undifferentiated Geneseo and Penn Yan shales)with a 25 mm (1 in) bed of fine-grained limestone, 0.20 m(8 in) above the base. There are small carbonate concentra-tions near the top of this interval, 76 mm (3 in) below theupper Genundewa Limestone. Bed 12/3, a 25 mm (1 in)-thick seam of laminated calcarenite on the underside of theupper Genundewa Limestone, contains the rich conodontand fish-bone fauna of the North Evans Limestone (see ref-erences under Loc. 13). A similar section on the western sideof the creek, below where the power lines cross NorthrupRoad, was illustrated by Brett & Baird (1982: 34, fig. 5B)and Baird & Brett (1986a: 166, fig. 4A). The section wasalso illustrated by de Witt & Colton (1978: pl. 2).

Locality 13*, Cayuga Creek.–The Cayuga Creek section,2 mi (3.2 km) northwest of Cowlesville (Wyoming County)shows the entry of some of the characteristics of more east-erly successions. The top of the illustrated section (Text-fig.8, Loc. 13) is 120 yd (110 m) upstream of the NY Rte. 354bridge, where the Genundewa Limestone forms a falls belowthe top of an old concrete dam. The interpretation of bedsdiffers in detail from accounts given earlier (see referencesbelow). As at Cazenovia Creek, the Leicester Pyrite forms abasal marker between the gray Moscow Shale and the blackGeneseo Shale. The 0.28 m (11 in)-thick black shale bandimmediately above the Leicester marks the base of the Gene-seo to the east (lower black shale tongue of de Witt &Colton, 1959: 2815). Beds 13/1 and 13/2 are thin blacklimestones in the succeeding 0.65 m (2.2 ft) of dark gray toblack shale with horizons rich in the diminutive brachiopodDevonochonetes and current-oriented Styliolina. Bed 13/2 oc-cupies the position of the Geneseo Limestone farther east.The 0.13 m (5 in)-thick black shale band above the intervalof dark gray to black shale is the upper black shale tongue(and top) of the Geneseo Shale. Above is a 10 mm (4 in)-thick interval of gray mudstone which is equivalent to theLodi Limestone horizon of the lower Penn Yan Shale to theeast. The overlying 0.23 m (9 in)-thick black shale is the

lower of the pair of black shales above the Lodi to the east(Penn Yan Black Shale A). It is not the Renwick Shale andtop of the Geneseo Shale as previously suggested (Kirch-gasser & Brett, in Oliver & Klapper, 1981b: 19, fig. 1; reil-lustrated by Kirchgasser, Brett, & Baird, 1997: 197, fig. 13).

Poorly preserved goniatites occur in Bed 13/3, theprominent bed of concretions in the Penn Yan, which is be-lieved to be the concretion horizon below the Schumacheror SB black shale farther east. Bed 13/4 is the Linden Hori-zon, a styliolinid-rich concretionary bed which farther eastis the source of well-preserved Koenenites, but no specimenshave been noted here. Bed 13/5 is the bed of grotesque con-cretions at the top of the Penn Yan Shale. Immediately abovethe concretion layer, mm- to cm-thick seams of North Evansfish-bone and conodont debris encrust the lower surface ofthe Genundewa Limestone. The reworked fish and con-odont debris of the North Evans, associated with the sub-upper Genundewa unconformity, occurs as far east asTaunton Gully (Loc. 17) in the Genesee Valley (Kirchgasser,1996a, 1998; Kirchgasser & Koslowski, 1996; Kirchgasser& Vargo, 1998). The section was illustrated by de Witt &Colton (1978: pl. 2). A detailed log was given by Brett &Baird (1990: A-19, fig. 6; reillustrated by Kirchgasser, Brett,& Baird, 1997: 195, fig. 12).

Locality 14*, Elevenmile Creek.–In Elevenmile Creek,the section illustrated commences approximately 230 yd(210 m) downstream of a railroad bridge, 1.3 mi (2.1 km)southwest of Darien Center (Genesee County). Bed 14/2, inthe Penn Yan Shale, 1.19 m (3.9 ft) below the upper Ge-nundewa, yields goniatites and correlates eastward with theLinden Horizon (Bed 15/8) at Linden (Loc. 15).

Locality 15*, Linden.–The section near Linden (GeneseeCounty) is in a small, eastward-flowing tributary to LittleTonawanda Creek, 0.3 mi (0.48 km) north of the bridge atLinden. The Moscow/Geneseo contact is at the confluenceof the tributary and creek. A group of black shales and con-cretion horizons is exposed in a falls section capped by theGenundewa Limestone (lower and upper parts). The LodiLimestone horizon is Bed 15/4, a bed of nodules beneath a0.61 m (2 ft)-thick black shale which is equivalent to thepair of marker black shales traceable above the Lodi eastwardto the Genesee Valley. Bed 15/5, the most prominent bed ofconcretions in the falls section, underlies a 0.30 m (1 ft)-thick black shale believed to be the SB marker black shale.Nearer the top of the Penn Yan Shale, the Linden Horizon(Bed 15/8) is expressed as a distinctive bed of nodules, some-times spherical and pyritic, with Koenenites beneath a 76mm (3 in)-thick seam of styliolinid limestone (Bed 15/9)which underlies a 0.18 m (7 in)-thick black shale, the baseof which is approximately 2.1 m (7 ft) below that of the Ge-nundewa Limestone. The Linden Horizon, which locally


yields well preserved K. styliophilus styliophilus, can be tracedeastward to the Genesee Valley and possibly as far east as theKeuka, Seneca, and Cayuga Lake valleys (Kirchgasser,1985). Shale partings at the base of Bed 15/9 yield crushedspecimens of Acanthoclymenia sp. The section was illustratedby de Witt & Colton (1978: pl. 2).

Locality 15a, Linden Falls.–A similar sequence is seen inthe face of the falls over the Genundewa, below the bridgeat Linden. There Bed 15a/5, the same horizon as Bed 15/5,lies 0.30 m (1 ft) above the level of the pool at the base ofthe falls. Bed 15a/8, the Koenenites-bearing Linden Horizon,occurs 1.96 m (6.4 ft) above Bed 15a/5.

Locality 16*, White Creek.–A discontinuous sectionthrough the Lower Genesee crops out along the westernbranch of White Creek near farm buildings downstream(north) of the NY Rte. 20 overpass, 2.1 mi (3.4 km) east ofBethany Center (Genesee County). In this section, theGeneseo Limestone horizon is the resistant 0.15 m (6 in)-thick limestone which caps a small falls downstream of thefarm, 3.5 m (11.5 ft) above the Moscow-Genesee contact.This is the westernmost locality where the Geneseo Lime-stone horizon has been identified in outcrop, but the hori-zon correlates westward into the cluster of threeconcretionary bands beneath the base of the upper blackshale tongue of the Geneseo as far west as Cayuga Creek(Loc. 13). Above the Geneseo Limestone Horizon, expo-sures are poor, but in an open field below the farm buildings,exposures resume with a horizon of large concretions (Bed16/3a) below the pair of marker black shales in the lowerPenn Yan Shale. Bed 16/3a is in the position of the LodiLimestone. Above this level, Ponticeras? occurs in the concre-tions of Bed 16/4 which lies 1.17m (3.8 ft) above the higherof the two black shales. A 51 mm (2 in)-thick black shale(?SB black shale) lies 0.61 m (2 ft) above. In the small em-bankment on the northern side of the creek, Bed 16/5 is thenodular Linden Horizon with Koenenites (thought to be thesame as Beds 15/8 and 15a/8 at Linden) and lies 0.13 m (5in) below the 51-mm-thick black shale. Bed 16/6 is a 76mm (3 in)-thick styliolinid band on top of Bed 16/5. Thingoniatite-bearing styliolinid limestones (Bed 16/6a, 16/7)are exposed near the culvert under NY Rte. 20. The higherbeds of the Penn Yan Shale are poorly exposed in the creekbed on the south side of NY Rte. 20, below outcrops of theGenundewa Limestone.

Locality 17*, Taunton Gully.–A well-exposed successionof the lower Genesee is found in Taunton Gully, approxi-mately 670 yd (612 m) upstream from the railroad crossingof Pearl Creek, 1.7 mi (2.7 km) north of Leicester (Liv-ingston County). Here the Geneseo Limestone (Bed 17/1)is the prominent 0.2 m (8 in)-thick unit 5 m (16.4 ft) abovethe Leicester Pyrite. The higher parts of the section are most

accessible near the two falls below the bridge where StarrRoad crosses the creek near the old Taunton Cemetery. TheLodi Limestone horizon is a distinctive horizon of smallnodules (Bed 17/2), 0.84 m (2.75 ft) above a 0.25 m (10in)-thick black shale exposed near the level of the large poolat the base of the lower falls; the top of the black shale marksthe Geneseo/Penn Yan contact. The Lodi bed locally yieldsPonticeras, and the overlying pair of black shales (the lowerone is Penn Yan Black Shale A, the upper Black Shale B)provide useful marker horizons on both sides of the GeneseeValley.

The 20-mm-thick SB marker black shale (Bed 17/2a)lies 0.38 m (1.25 ft) above the first prominent bed of con-cretions (with indeterminate goniatites) in the face of themain falls and 3.8 m (12.6 ft) above the Lodi horizon. Koe-nenites occurs in Bed 17/3, a horizon of concretions 0.58 m(1.9 ft) above the SB black shale and 1.2 m (4 ft) below astill higher concretionary horizon (Bed 17/4) with Koenen-ites and Epitornoceras? at the lip of the falls. The main Koe-nenites horizons (K. styliophilus styliophilus) occur above thelip of the falls in the concretions of Bed 17/5 and immedi-ately above in the pyritic nodules of the Linden Horizon(Bed 17/6). The Linden underlies a 0.13 m (5 in)-thickblack shale and is 5.03 m (16.5 ft) below the base of the Ge-nundewa Limestone, which crops out below the bridge. Thesection was illustrated by deWitt & Colton (1978: pls 2, 6).

Locality 17a, Spezzano Gully.–A section similar to thatin the upper part of Taunton Gully is exposed in SpezzanoGully west of the railroad crossing, 0.9 mi (1.4 km) farthernorth, and 2.6 mi (4.2 km) north of Leicester (LivingstonCounty). The productive horizons are Bed 17a/3, a horizonof large concretions 0.91 m (3 ft) above the lip of the mainfalls, and Bed 17a/4, which is taken as the main Koeneniteshorizon (Linden Horizon), a layer of nodules on top of a0.13 m (5 in)-thick limestone and beneath a 76 mm (3 in)-thick black shale, 0.96 m (3.2 ft) above Bed 17/a3 and 4.1m (13.5 ft) below the Genundewa Limestone.

Locality 18*, Beards Creek.–Along Beards Creek, 0.3 mi(0.48 km) northwest of Leicester (Livingston County), is deWitt & Colton’s (1959) reference section for the GeneseeGroup. The section commences above the Leicester Pyrite,0.1 mi (0.16 km) from the railroad crossing of the creek.Here a 1.2 m (4 ft)-thick, lower black shale tongue of theGeneseo Shale is followed by 6.8 m (22.4 ft) of dark grayshale with thin limestones that in turn are overlain by a 0.38m (1.25 ft)-thick upper black shale band. The two blackshale units respectively represent deWitt & Colton’s (1959)lower and upper tongues of the Geneseo. The interveningdark gray shales were regarded by them as a tongue of PennYan Shale, but they are now included in the Geneseo Shale.Tornoceras is relatively common in Bed 18/1, within the first


0.3 m (1 ft) above the top of the lower tongue of black shale.Bed 18/2 lies immediately above the highest of five thinblack shales in the succeeding 1.8 m (5.8 ft) of the section.

The Geneseo Limestone (Bed 18/2a), the prominentlimestone in approximately the middle of the gray shale in-terval, lies 3.9 m (12.8 ft) above the top of the lower blackshale tongue. Bed 18/3, also with Ponticeras, is a thin lime-stone 0.35 m (1.2 ft) above. The nodular Lodi Limestone(Bed 18/4) with P. perlatum lies beneath a 76 mm (3 in)-thick black shale (Black Shale A of the Penn Yan Shale), 1.42m (4.6 ft) above the top of the upper black shale tongue ofthe Geneseo Shale and 0.66 m (2.2 ft) above the top of the0.3 m (1 ft)-thick black shale, the top of which defines thecontact between Geneseo and Penn Yan Shales. Ponticerasalso occurs in an interval of gray shale (Bed 18/5) exposedin the southern side of the creek, downstream of the highfalls over the Genundewa Limestone below the DunkleyRoad bridge. Bed 18/5 is the interval 1.7-4 m (5.6-13.2 ft)above the Lodi, and the large septarian nodules (Bed 18/6)just above might be the concretion horizon below the SBblack shale. Horizons higher than Bed 18/6 are not readilyaccessible for collecting. The section was illustrated by deWitt & Colton (1978: pl. 5).

Locality 19*, Fall Brook-Dewey Hill.–On the easternside of the Genesee Valley, the well-known locality noted byJames Hall (1843: 462) at Fall Brook, 1.5 mi (2.4 km) southof Geneseo (Livingston County), provides a spectacular ex-posure from the upper Moscow Shale through the lowerGenesee Group. In the cliff above the northern bank of thecreek downstream of the falls over the Genundewa Lime-stone, the lower and upper tongues of the Geneseo Shale,each approximately 1.5 m (5 ft) thick, are separated by 5.2m (17 ft) of dark gray shale. Lenses of the Leicester Pyrite arewell exposed at the Moscow-Geneseo contact. The promi-nent 0.17 m (7 in)-thick limestone (Bed 19/3) with crushedPonticeras, in approximately the middle of the gray shale in-terval, is the Geneseo Limestone. This widespread horizonin the Genesee Valley, which there lies 4 m (13 ft) above thebase of the lower black shale tongue, was previously referredto the Lodi Limestone (Kirchgasser, 1975; Kirchgasser &House, 1981; Oliver & Klapper, 1981b), but that horizonis now known to be higher in the section.

The section is accessible in a small tributary that entersFall Brook at the eastern edge of the cliff. In the side creek,the 1.4 m (4.5 ft)-thick upper black shale tongue of theGeneseo Shale lies 0.20 m (8 in) above a thin limestone (Bed19/3a), which caps a falls. The Lodi Limestone is the nodu-lar bed (Bed 19/4) in the lower Penn Yan Shale that lies be-neath a thin black shale (Black Shale A), 2.9 m (9.5 ft) abovethe base of the upper black shale tongue of the GeneseoShale.

The higher part of the Penn Yan can be measured in theside creek, but a more accessible continuation of the succes-sion is seen in the NY Rte. 39-20A roadcut along DeweyHill, 0.2 mi (0.32 km) to the north. The nodules of Bed19/4 (Lodi Limestone) and the capping 77 mm (3 in)-thickblack shale (Black Shale A) are exposed in the ditch on thesouthern side of the road where the outcrop begins. Theprominent 0.18 m (7 in)-thick black shale (Bed 19/4a),overlying a line of concretions, approximately 5.2 m (17.2ft) above the Lodi, is the SB black shale horizon. Althoughnot yet recognized in the Genesee Valley sections, the blackRenwick Shale and the succeeding Abbey Gulf Beds of moreeastern sections are believed to project into the interval be-tween the Lodi and SB horizons.

In the gray shales 1.2 m (4 ft) above the SB horizon is aprominent layer of large concretions (Bed 19/4b), and 1.8m (5.75 ft) above is Bed 19/5, the first of a series of thinlimestones with goniatites. Preservation is poor but expo-sures of Bed 19/5 on the northern side of the road haveyielded Ponticeras? and a large tornoceratid (?Epitornoceras).Bed 19/5 lies 8.6 m (28.25 ft) above the Lodi horizon. Onthe southern side of the road, Bed 19/6, which lies 0.97 m(3.2 ft) above the base of Bed 19/5, is the Linden Horizon,the first of a series of thin styliolinid beds. Bed 19/6 containssmall baritic replacements of Koenenites styliophilus stylio-philus and Tornoceras. Goniatites also occur in the Bed 19/7approximately 0.42 m (1.4 ft) above the base of Bed 19/6and in the highest styliolinid band (Bed 19/8), which is 3.07m (10 ft) above the base of Bed 19/6 and approximately 3m (10 ft) below the base of the Genundewa Limestone,which crops out in the bushes at the top of the section.

This section has been illustrated by de Witt & Colton(1978: pl. 2), Oliver & Klapper (1981b: 37), Kirchgasser etal. (1994: 51), and Brett & Ver Straeten (1997: 222).

Localities 20* and 20a, Hemlock.–The nearest useful ex-posures of the lower Genesee east of Fall Brook are thosenorth of Hemlock (Livingston County), almost 13 mi (21km) distant, a fact that makes detailed correlation difficult.In the westward-flowing tributary to Hemlock Outlet,which crosses Big Tree Road 0.9 mi (1.4 km) northeast ofHemlock (Loc. 20), the upper part of the Penn Yan Shale iswell exposed below a falls above the Genundewa Limestone.A a similar setting is found in another tributary (Loc. 20a)that crosses the Huff-Barnard road intersection 0.9 mi (1.4km) farther north. Beds 20/1, 20/2, and 20/3 are thin lime-stones with goniatites that crop out within the first 2.7 m (9ft) above where exposures begin. At Loc. 20a, thin blackshales were noted above equivalents of Beds 20/2 and 20/3.In the upper part of the main section, Beds 20/4 and 20/5form a pair of thin limestones (with poor goniatites) 50-75mm (2-3 in) thick and 0.3 m (1 ft) apart. These occur 1.75


m (5.75 ft) below a layer of large concretions at the foot ofthe falls below the Genundewa Limestone. Bed 20/4 (and20a/4) is taken as the Linden Horizon.

Locality 21, Abbey Gulf.–A well-exposed and almostcomplete section through the lower Genesee Formation isfound in Abbey Gulf, east of Allens Hills Road, 1.6 mi (2.6km) northeast of Honeoye (Ontario County; Baird et al.,1989: figs. 7A, 9B). Although the goniatite record there ispoor, several marker beds in the section are important forcorrelations eastward to Canandaigua Lake and beyond.References are made to the sections by de Witt & Colton(1978: pl. 2): Cd-Abbey Gulf, Cd-6-Vincent, and Cd-7-Menteth Gully (Canandaigua Lake). At Abbey Gulf andnearby sections in the Honeoye, Bristol, and CanandaiguaLake valleys, the Middle-Upper Devonian (Givetian-Frasn-ian) boundary is located above the Lodi Limestone in thelowermost Penn Yan Shale (Kirchgasser et al., 1989; Kirch-gasser, 1994: fig. 3).

In the lower part of the Abbey Gulf section, at a leveltaken to be the top of the upper black shale tongue and topof the Geneseo Shale, is the Lodi Limestone (Bed 21/1), a0.1 m (4 in)-thick fossiliferous limestone with auloporidcorals, brachiopods, and crinoid fragments. The Lodi is cutout in the main creek section but the horizon lies 0.91 m (3ft) above the first low falls in the section. The Lodi is ex-posed in a tributary, 4.1m (13.4 ft) above where the tribu-tary enters from the north. Above the position of the Lodihorizon in the main creek section, a 2 m (6.5 ft)-thick inter-val of olive gray shale and mudrock is overlain by a 0.61 m(2 ft)-thick black shale at the top of a low falls. A Ponticeraswas noted in Bed 21/1c at the erosional base of the blackshale. The black shale is Penn Yan Black Shale B, the higherof the pair of black shales above the Lodi in the GeneseeValley (Text-fig. 8), and the first thick black shale above theLodi in the Menteth Gully section (Cd-7) at CanandaiguaLake (de Witt & Colton, 1978: pls 2-3).

Above Penn Yan Black Shale B in Abbey Gulf is a 2.7 m(9 ft)-thick interval of siltstone and silty shale (a tongue ofSherburne Siltstone), at the top of which is a 0.53 m (1.75ft)-thick black shale taken to be the Renwick Shale, a unittraceable to Canadaigua Lake (Loc. Cd-7) and eastward toits type area near Cayuga Lake where it lies between theSherburne Siltstone and the Ithaca Shale and Sandstone (deWitt & Colton, 1978: pls 2-3).

Above the Renwick Shale, poor goniatites occur in Beds21/1a and 21/1b below and above a 0.30 m (1 ft)-thickblack shale, within a 2.6 m (8.5 ft)-thick interval of irregu-larly bedded silty shales and siltstones (some with barite andpoor goniatites); these are the informal “Abbey Beds” of G.Baird and C. Brett (Baird et al., 2006: 355; Zambito et al.,2007: 94, fig. 4). The “Abbey Beds” are capped by a 0.30 (6

in)-thick black shale taken to be the SB black shale of west-ern sections, which thus constrains the interval of the Sher-burne, Renwick, and “Abbey Beds” to the interval betweenthe Lodi and the SB black shale horizon in the Genesee Val-ley and farther westward (Text-fig. 8). The SB black shale isthe first black shale below sample-horizon 6697 SD inMenteth Gully (de Witt & Colton, 1978: pls 2-3).

Approximately 3 m (10 ft) above the SB black shale, a0.10 m (4 in)-thick limestone, beneath a 0.61 m (2 ft)-thickblack shale, caps a low falls. This limestone correlates west-ward with Bed 20/3 at Hemlock. Cropping out 2.4 m (8 ft)above the top of the falls is a pair of thin limestones, Bed21/2, 0.10 m (4 in) thick, and Bed 21/3, 76 mm (3 in)thick, 0.30 m (1 ft) higher. Although no goniatites havebeen found, Beds 21/2 and 21/3 are taken to be the LindenHorizon, which correlates westward to Beds 20/4 and 20/5at Hemlock and Bed 19/6 at Fall Brook-Dewey Hill (Text-fig. 8). At Abbey Gulf, a gas seep reaches the surface in thebeds of the upper Penn Yan below the falls capped by theGenundewa Limestone. Bed 21/5 is a 76 mm (3 in)-thickbed of nodules with Acanthoclymenia and Tornoceras that lieswithin black shale, 0.38 m (1.25 ft) below the Genundewa.

Locality 22, Barnes Gully.–A long and complete sectionof the lower Genesee is exposed in Barnes Gully, 2.5 mi (0.4km) southeast of Cheshire (Ontario County), on the west-ern side of Canandaigua Lake, above Foster Point. The sec-tion commences above the Windom/MoscowShale-Geneseo Shale contact at the top of a falls at approx-imately 800 ft (244 m) altitude. The section illustrates thegreat eastward thickening of the Geneseo Shale. No goni-atites were collected at this locality.

Locality 23*, Seneca Point Creek.–A section in the upperPenn Yan Shale is exposed below the first falls in SenecaPoint Creek above Seneca Point on the western side ofCanandaigua Lake. Bed 23/a, a bed of concretions 7.1 m(23.3 ft) below the base of the Genundewa Limestone, isthe “hiatus-concretion” horizon of Baird (1976). The hori-zon lies below the level of the pool at the base of the falls,0.61 m (2 ft) above a pair of black shales, 0.38 m (1.25 ft)and 76 mm (3 in) thick. Bed 23/a contains abundant andwell-preserved pyritic and baritic protoconchs and earlywhorls of Koenenites and Tornoceras and a distinctive con-odont fauna with Ancyrodella crosbiensis Kralick, 1994, indi-cating a correlation with the Crosby Sandstone of Torrey etal. (1932) to the east and possibly also with the LindenHorizon to the west; however, the Linden Horizon mightbe lower in the succession. Bed 23/a and the underlyingblack shales were illustrated by deWitt & Colton (1978: pls2-4) in the Seneca Creek section (Nap-1). The lower andthicker of the black shales could be the “Starkey” black shaleof de Witt & Colton (1978: pls 4-5).


Locality A, Nap-4, Genundewa Point.–On the easternside of Canandaigua Lake, north of Genundewa Point, theequivalent of Bed 23/a of Seneca Point Creek is Bed 23/b,a 0.23 m (9 in)-thick concretion bed, beneath a prominent0.20 m (8 in)-thick siltstone, 5.9 m (19.25 ft) below thebase of the Genundewa Limestone. The horizon and cap-ping siltstone were illustrated by de Witt & Colton (1978:pl. 4).

Locality B, Ph-5, Shuman Cemetery.–In the tributary ofFlint Creek 0.8 mi (1.3 km) north of Shuman Cemeteryand 2 mi (3.2 km) south of Gorham (Ontario County), theGenundewa Limestone, cropping out at approximately 936ft (285 m) elevation, has thinned to a single prominent styli-olinid limestone several centimeters thick, above black styli-olinid-rich shales. In the upper Penn Yan Shale, 1.2 m (4 ft)below the Genundewa, Gordon Baird collected a 10 mm (4in)-thick styliolinid bed with numerous baritic Koenenitesstyliophilus. The horizon is in a covered interval and mustbe dug out to be collected. The bed could represent eitherthe Linden Horizon or the Crosby Sandstone. At the top ofthe Genundewa at this locality, Gordon Baird (pers. comm.,2002) has found pyritic goniatites in a 1-3 cm-thick corro-sional lag marking a discontinuity traceable southeastwardto Sunset Point ravine on Keuka Lake (see below). This sec-tion was illustrated by de Witt & Colton (1978: pl. 4).

Locality C, Py-11, Sartwell Ravine.–The upper PennYan Shale and lower Ithaca Shale and Sandstone are exposedin Sartwell Ravine in the vicinity of the Kimble Road bridgecrossing, beside the airport, 1 mi (1.6 km) south of PennYan (Yates County). The Crosby Sandstone, at or near thePenn Yan-Ithaca contact, is the source of a large collectionof Koenenites styliophilus styliophilus made by D. D. Lutherin 1904 (his locality 3397) found in the NYSM (Kirch-gasser, 1982, 1985). The position of the Crosby Sandstonein the section is uncertain. Luther’s label with the collectionnotes the horizon at 880 ft (268.2 m) elevation, whichwould be approximately 200 yd (183 m) above the KimbleRoad crossing. The Crosby was not found in place in asearch in 1977 of the several hundred yards (meters) of in-termittent exposures of the Ithaca upstream of the bridge inthe tributary paralleling the new NY Rte. 14A. Bergin(1964: 11) found the Penn Yan-Crosby contact to be wellexposed at an altitude of 860 ft (262.1 m), although on hismap the contact was drawn at approximately 870 ft (265m), which is at approximately the level of the Kimble Roadbridge and within 10 ft (3 m) of Luther’s locality. De Witt& Colton (1978: pl. 4, table 1) placed the base of theCrosby at 811 ft (247.2 m) elevation, 5.8 m (19 ft) abovethe base of the Renwick Shale, or approximately 21 m (70ft) below Luther’s locality

Bed Py-11/a consists of loose blocks of goniatite-rich

Crosby Sandstone found approximately 183 m (600 ft)downstream of the Kimble Road bridge. Bed Py-11/b is aloose block of Crosby with goniatites from approximately110 m (360 ft) upstream of the bridge; this bed number isused for Luther’s collection. Bed Py-11/c is a loose block ofcalcareous siltstone with goniatite fragments from approxi-mately 183 m (600 ft) upstream of the second of two dirttracks which cross the creek approximately 366 m (0.23 mi)above the bridge.

Locality D, Py-11a, Lakeside Country Club.–Blocks ofquestionable Crosby Sandstone were found in the creek onthe northern side of the Lakeside Country Golf Club, 1.7 mi(2.7 km) southwest of Penn Yan (Yates County), and aboveKeuka Lake (East Branch). The locality is below the seventhgreen at approximately 800 ft (244 m) elevation and approx-imately 622 m (0.39 mi) west of the Bath Road (NY Rte.17) and near the Penn Yan-Crosby contact as mapped byBergin (1964).

Locality E, Py-14,Willow Grove.–The uppermost PennYan Shale, the Crosby Sandstone, and lower tongue of theIthaca Shale and Sandstone are exposed in a waterfall sectioninWillow Grove Gully that crosses NY Rte. 54, 3.5 mi (5.6km) southwest of Penn Yan (Yates County), above KeukaLake (East Branch). The sections begin at approximately780 ft (238 m) elevation, 0.3 mi (0.48 km) east of the NYRte. 54 bridge. Bed Py-14/1 is the goniatite-rich basal 10cm (4 in)-thick unit of the 2 m (6.6 ft)-thick Crosby Sand-stone, a calcareous siltstone with reworked concretions; in-cluded is Tornoceras arcuatum. This section was illustrated byde Witt & Colton (1978: pl. 4).

Locality F, Py-16, Sunset Point Gully.–The upper PennYan Shale, the Crosby Sandstone, and lower tongue of theIthaca Shale and Sandstone are well exposed in Sunset PointGully, above Sunset Point, Keuka Lake (East Branch) thatcrosses NY Rte. 54, 0.2 mi (0.32 km) south of Sisson Roadand approximately 5.8 mi (9.3 km) southwest of Penn Yan(Yates County). Exposures begin in the upper Penn Yan ap-proximately 15 m (49 ft) east of NY Rte. 54. The 20 cm (8in)-thick Crosby Sandstone caps the first high waterfall andthe Ithaca beds are well exposed above.

Koenenites styliophilus styliophilus is common in the go-niatite fauna at the base of Bed Py-16/1, the Crosby Sand-stone, a level that also yields Tornoceras arcuatum. A finepyritic specimen of Acanthoclymenia sp. was collected byJames Kralick from a concretion in the uppermost Penn YanShale 0.30 m (1 ft) below the base of the Crosby Sandstone.Additional goniatites from the Crosby Sandstone were col-lected from loose blocks on the creek floor approximately69 m (226 ft) east of NY Rte. 54. This section was illustratedby de Witt & Colton (1978: pl. 4).

Locality G, Ov-3a, Gully north of Plum Point, Seneca


Lake.–In an unnamed creek immediately north of PlumPoint Road, north of Plum Point, Seneca Lake, 1.3 mi (2.1km) northeast of Himrod (Yates County), poor Ponticerasoccur in a dark silty shale above lenses of pyrite equivalentto the Lodi Limestone (Lodi Bed A) at the Geneseo Shale-Penn Yan Shale contact. This is locality 12B of Baird et al.(1989).

Locality H, Ov-4, north of Baskin Point, SenecaLake.–On the northern side of Baskin Point, Seneca Lake,1.4 mi (2.3 km) northeast of Starkey (Yates County), boththe Lodi Limestone (Lodi Bed B; Bed 25a/1) and the over-lying massive siltstone (Bed 25a/2) contain goniatites in a se-quence similar to that seen to the south at Fir Tree Point(Loc. 25) and across the lake at Lodi Glen (Loc. 27).

Locality I, Ov-5, south of Baskin Point, Seneca Lake.–Inthe creek 0.2 mi (0.32 km) south of Baskin Point, SenecaLake, a single specimen of Ponticeras was recovered in one ofthe upper cycles of the Lodi Limestone, approximately 5.2m (17 ft) above the Lodi horizon at the Geneseo Shale-PennYan Shale contact (sample collected by WTK and GordonBaird).

Locality J, Wg-2, north side of Fir Tree Point.–In theembankment, creek, and access road (Rock Stream Road)on the northern side of Fir Tree Point, Seneca Lake, poorgoniatites occur in the styliolinid and ostracod-rich blackshale capping the Fir Tree Pyrite Bed. Above the Fir TreeBed is a 0.68 m (2.25 ft)-thick interval of black shale at thetop of the Geneseo Shale (Hubbard Quarry Shale) withinwhich is a nodular horizon (informal “Bergen Beach Bed” ofBrett & Baird, 1986). Above is the first of four calcareousbeds comprising the Lodi Limestone (Lodi Beds A, B, C,and D) through an interval of 4.3 m (14 ft). The 0.71 m(2.3 ft)-thick main bed of the Lodi Limestone (Lodi bed B)crops out 1 m (3.3 ft) above the Fir Tree bed. No goniatiteswere noted in the Lodi interval at this locality. This islocalityWg-2 of deWitt & Colton (1978) and locality 16Aof Baird et al. (1989).

Locality 25, Wg-3, Fir Tree Point.–On the western sideof Seneca Lake, 1.2 mi (1.9 km) southeast of Rock Stream(Yates County), at the southern side of Fir Tree Point (for-merly Rock Stream Landing), the top 1.8 m (6 ft) of theGeneseo Shale is exposed (Loc. 25). Above this is a blocky-weathering massive calcareous siltstone approximately 1.8m (6 ft) in thickness (Bed 25/2) with a 0.3 m (1 ft) moreconcretionary unit, below (Bed 25/1) (probably Lodi Lime-stone Bed B) and a shale unit above. Locality Wg-3 was il-lustrated by de Witt & Colton (1978: pl. 5).

Locality K, Wg-7a, Green Point, Seneca Lake.–Theupper Geneseo Shale, Lodi Limestone, and lower Penn YanShale are exposed in the bluffs and roadcuts behind cottagesalong the shore of Seneca Lake between the second and third

unnamed points southeast of Fir Tree Point, approximately1.8 mi (2.9 km) southeast of Rock Stream (Yates County).The access road begins 0.1 mi (0.16 km) south of the inter-section of the Bates Cross Road and NY Rte. 14. The bedsof the Lodi Limestone (Lodi Beds A, B, C, and D) are ex-posed through a 2.9 m (9.5 ft)-thick interval and, higher inthe Penn Yan Shale, Ponticeras occurs in a 0.18 m (7 in)-thick calcareous siltstone with a fauna of large styliolines,gastropods, and brachiopods that lies 2.4 m (8 ft) above thetop of the Lodi and immediately below a 1.2 m (4 ft)-thickblack shale (locality 16B of Baird et al., 1989).

Locality L, Ov-10, Mill Creek.–The equivalents of theupper Penn Yan Shale and Crosby Sandstone occur in atongue of the Ithaca Shale and Sandstone exposed in MillCreek, below the Upper Lake Road bridge, 1.3 mi (2.1 km)west of Lodi (Seneca County; eastern side of Seneca Lake).The bridge foundation overlies the massive 0.69 m (2.25ft)-thick Crosby Sandstone, and approximately 17 m (56 ft)of Penn Yan Shale equivalents are well exposed in the creekbelow.

Bed Ov-10/1, with a few well preserved Koenenites stylio-philus styliophilus, is believed to be the Linden Horizon. It isa nodular band with nautiloids, brachiopods, and auloporidswithin a silty shale and mudrock interval that overlies an in-terval of siltstones (some with ripplemarks) that cap a seriesof falls; the horizon is 6.1 m (20 ft) below the base of theCrosby Sandstone. Bed Ov-10/2 at the top of the silty shaleand mudrock interval is a 0.28 m (11 in)-thick, irregularlybedded, silty mudrock with brachiopods, tabulates, and bi-valves and a goniatite fragment with the growth lines of?Chutoceras nundaium (Hall, 1874). The top of the bed is4.6 m (15 ft) below the base of the Crosby Sandstone.Poorly preserved Koenenites were noted in Bed Ov-10/3 inthe float immediately beneath a prominent 0.18 m (7 in)-thick siltstone, the top of which is 3.2 m (10.5 ft) below thebase of the Crosby Sandstone. Acanthoclymenia occurs inBed Ov-10/4, a silty mudrock, 2.3 m (7.5 ft) below the baseof the Crosby Sandstone. The horizon lies below a pair ofthin siltstones and is 0.91 m (3 ft) below the top of the 0.15m (6 in)-thick ripple-marked siltstone that caps a low falls.Above the ripple-marked siltstone, Bed Ov-10/5 is a 76 mm(3 in)-thick mudrock with a shelly fauna and poorly pre-served Koenenites? that lies 0.91 m (3 ft) below the base ofthe Crosby Sandstone. This section was illustrated by deWitt & Colton (1978: pl. 3) and Kirchgasser (1981: 45).

Locality M, Ov-13, north of Tommy Creek.–Large butpoorly preserved specimens of Ponticeras cf. perlatum, andrare pyritic goniatite protoconchs in conodont residues,occur in a 9 cm (4 in)-thick bed of Lodi Limestone (LodiBed C) collected by Gordon Baird at a level 5.5 m (18 ft)above the main Lodi Limestone horizon (Lodi B) in the


gully 0.3 mi (0.48 km) north of Tommy Creek, 2.6 mi (4.2km) northwest of Lodi (Seneca County). The main LodiLimestone bed (Lodi B) with a pyritic crust at its top, is atthe top of a waterfall 15 ft (4.6 m) high, and approximately0.30 m (1 ft) above the top of the Geneseo Shale (HubbardQuarry Shale). Pyritic goniatite protoconchs were noted inthe conodont residues from the top of Lodi Bed B. DeWitt& Colton (1978: table 1) placed the top of the Geneseo at638 ft (194.5 m) elevation. This is locality 18 of Baird et al.(1989).

Locality 27, Lodi Glen.–Above the Tully Limestone inLodi Glen (Loc. 3), the Geneseo Shale and Penn Yan Shaleare exposed in a falls section below the railroad bridge, 1.9mi (3.0 km) west of Lodi (Seneca County). An interval ofconcretions in the lower Penn Yan Shale (or Sherburne Silt-stone) is the type section of the Lodi Limestone, and is thesource of Ponticeras perlatum, which also occurs, crushed flat,in the shales below. Bed 27/1, the main Lodi bed (Lodi B),with P. perlatum, is a 0.23 m (9 in)-thick concretionaryband, the base of which is 1.2 m (4 ft) above the top of theGeneseo Shale (Hubbard Quarry Shale); this is locality 3013of Kirchgasser (1975). Pyritic goniatite protoconchs andshell fragments are common in the conodont-bone-bed hashcomprising Lodi Bed A at the contact between the Geneseoand Penn Yan Shales. A representative of the Epitornocerasperacutum group (E. cf. mithracoides) was collected by J. W.Wells at this locality from a level 12.2 m (40 ft) below thetop of the Geneseo Shale. Section Ov-10 was illustrated byde Witt & Colton (1978: pl. 3), Oliver & Klapper (1981b:45, fig. 1), and Grasso et al. (1986: 174, fig. 4). This is lo-cality 17 of Baird et al. (1989) and Kirchgasser (1994).

Locality 27a; Ov-16, Romulus Town Quarry.–There areother sections between Seneca Lake and Cayuga Lake inwhich this same interval can be seen. In the large shale pitof the RomulusTown (Ovid) Quarry (Loc. 27a), 1.1 mi (1.8km) northwest of Ovid (Seneca County), approximately300 m (984 ft) along a track leading south fromWest BlaineRoad, halfway between NY Rte. 96 and the Kinne Road,the Lodi Limestone (Lodi B, Bed 27a/1) and associated lev-els occur with the typical Ponticeras perlatum fauna. This islocality 3012 of Kirchgasser (1975) and locality 19 of Bairdet al. (1989). This section was illustrated by Baird & Brett(1986b: 48, fig. 9F; 64, fig. 13) and Baird et al. (1989: 364,fig. 7F).

Locality 27b; Gen-2, Hubbard Quarry.–The abandonedshale pit of Hubbard Quarry (Loc. 27b) is immediately eastof NY Rte. 98 and just north of Lively Run, 0.12 mi (0.2km) south of the Ovid/Romulus town line and 1 mi (1.6km) southwest of Kidders (Seneca County). There the upperpart of the black Geneseo Shale (Hubbard Quarry Shaletype section) and lower Penn Yan Shale (Sherburne of au-

thors) crops out and the main Lodi Limestone horizon (LodiB) is a fossiliferous calcareous siltstone with auloporid coralsnear the base of the Penn Yan. Ponticeras perlatum occurs inthe topmost beds of the Geneseo Shale and in the Lodi bed;many specimens from both horizons are housed in the PRIcollections. This section was illustrated by Cole et al. (1959),deWitt & Colton (1978: pl. 3), Grasso et al. (1986: fig. 10),Baird & Brett (1986b: 41, fig. 5A; 64, fig. 13), andThomp-son & Newton (1987: fig. 3). This is locality 22 of Baird etal. (1989) and Kirchgasser (1994).

Locality P, Gen-1a, Ovid-Sheldrake Road.–A specimenof Ponticeras perlatum (CU 41083) was collected by J. W.Wells from an outcrop at an elevation of 720 ft (219.5 m)on the Ovid-Sheldrake (Parish) Road, approximately 0.7 mi(1.1 km) east of Sheldrake Springs (Seneca County). Wells’slocality 204, below Hall Road, where the creek parallel toand north of Sheldrake Creek crosses the Ovid-SheldrakeRoad, is probably in the Lodi Limestone of the lower Sher-burne Siltstone. The locality was not visited in this study.

Locality Q, I-1, Glenwood Creek.–In Glenwood Creekabove Glenwood Point, Cayuga Lake, approximately 5 mi (8km) south of Taughannock Point, Kindle (1896) collectedspecimens referable to Ponticeras perlatum from the upperSherburne Siltstone, at his locality 8-1, 12 m (40 ft) belowthe second “Spirifer laevis Zone” (CU 10934b; G125) andfrom the second “Spirifer laevis Zone” at the base of the over-lying Renwick Shale, his locality 8-2 (CU 10916; G137-138).

De Witt & Colton (1978: table 1, pl. 3) recorded thebase of the Renwick Shale (second “Spirifer laevis Zone” andKindle’s locality 8-2) at 634 ft (193.2 m) elevation, whichwould be upstream of the railroad crossing of the creek. Thesecond “Spirifer laevis Zone” refers to a bed or beds with thebrachiopod Warrenella laevis (Hall, 1843), which earlyworkers used to define the boundary between the SherborneSiltstone and the Ithaca Shale and Sandstone. Kindle’s Loc.8-1 in the underlying Sherburne Siltstone would be at 594ft (181.0 m) elevation, above NY Rte. 89 near where therailroad track crosses the creek.

The section continues above the Renwick Shale into theIthaca Shale and Sandstone. De Witt & Colton (1978)recorded the top of the Crosby Sandstone at a position 47m (155 ft) above the base of the Renwick at 793 ft (241.7m) elevation (downstream of the Duboise Road crossing).The section was not examined in this study.

Locality R, I-2, Williams Brook.–A section through theRenwick Shale and lower and middle Ithaca Shale and Sand-stone is exposed in Williams Brook, a creek on the south-western side of Cayuga Lake above the site of the oldmunicipal airport on the northwestern side of Ithaca (Tomp-kins County). Downstream of the NY Rte. 96 bridge is the


3-m-thick Williams Brook Coquinite of Caster (1933), thetop of which, at 620 ft (189.0 m) elevation, is approximately53 m (175 ft) above the base of the Renwick Shale (at 445ft or 135.6 m elevation) (de Witt & Colton, 1978: A-16,table 1). Two specimens in the CU collection referred herewith question to Chutoceras nundaium might have comefrom the Williams Brook Coquinite. The specimens (CU10928 and 10938) were collected by Kindle (1896) fromhis locality 4-5, a bed of impure limestone 265 ft (80.7 m)above the lake. Given a lake level of 382 ft (116.4 m) eleva-tion, that would place theWilliams Brook horizon at 647 ft(197.2 m) elevation, approximately 53 m (175 ft) above theRenwick Shale, a position close to but higher than thatrecorded by de Witt & Colton (1978). De Witt & Colton(1978: A-16) placed theWilliams Brook Coquinite approx-imately 21-24 m (70-80 ft) below the Crosby Sandstone.No new goniatites were found at this locality.

Locality S, I-2a, south of Williams Brook.–At a quarrynear the railroad approximately 0.5 mi (0.8 km) south ofWilliams Brook, Kindle (1896) collected a goniatite frag-ment here assigned with question to Chutoceras nundaiumbased on its growth lines and flattened venter. The specimencame from Kindle’s locality 4-2, 4.6 m (15 ft) above the sec-ond “Spirifer laevis Zone,” a bed or beds with the brachio-pod Warrenella laevis. The horizon is at or near the base ofthe black Renwick Shale.

Kindle’s locality 4-2 in the vinicity of the base of theRenwick Shale must be near H. S. Williams’ (1884) Station4 locality “at the southwest corner of Cayuga Lake” and pre-sumably near the Spirifer laevis beds at the top of the Sher-burne Siltstone and base of the Renwick Shale. Williams’Station 4 yielded specimens referable to Ponticeras perlatum(CU 41078-HSW coll. 4A, 41075-HSW Coll. 4B) and aninvolute specimen (HSW 4-D35; Cornell locality label, G-62) that might be a Koenenites. No new goniatites werefound at this locality.

Locality T, I-11d, Gulf Creek.–From the upper Sher-burne Siltstone in Gulf Creek (Shurger or Shurger’s Glen)above Portland Point, approximately 1 mi (1.6 km) south-east of South Lansing (Tompkins County), R. P. Boeken-kamp collected a poor specimen of Ponticeras perlatum (CU41074) at an elevation of 856 ft (260.9 m). The horizonwould be near the NY Rte. 34 crossing of the creek. On afield map produced by J. W.Wells, the top of the SherburneSiltstone (second “Spirifer laevis Zone” of authors) is at ap-proximately 940 ft (286 m) elevation, just upstream of theNorth Triphammer Road crossing, which would place thehorizon of CU 41074 approximately 26 m (84 ft) below thetop of the Sherburne. The section was not examined in thisstudy.

Locality U, I-11b, Esty Glen.–In Esty or Esty’s Glen,

above Esty Point at the southeastern end of Cayuga Lake,approximately 1.3 mi (2.1 km) north of McKinneys (Tomp-kins County), the upper Sherburne is exposed below NYRte. 34. The top of the Sherburne indicated by the second“Spirifer laevis Zone” of authors (beds withWarrenella laevis)has been given as either at 637 ft (194 m) (Williams, 1951)or 616 ft (187.8 m) elevation (Boekenkamp, 1963). Froman unknown level in the Sherburne, F. B. Hine collected (in1876) a Ponticeras perlatum (CU 41079). The section wasnot examined in this study.

Locality V, I-11a, Twin Glens at McKinneys.–The upperpart of the Sherburne Siltstone is exposed in Twin Glens,above McKinneys Point at the southeastern end of CayugaLake, just north of McKinneys or McKinney’s Station(Tompkins County). In the northern glen, the second “Spir-ifer laevis Zone” of authors, which marks the top of the Sher-burne Siltstone (and base of Renwick Shale), is atapproximately 502 ft (153 m) elevation (Boekenkamp,1963). Kindle (1896) collected specimens of Ponticeras per-latum (CU 19030a-b) from his locality 7-4, from the first S.laevis horizon of the Sherburne, which is approximately 33.5m (110 ft) below the top of the Sherburne and 20 ft (6.1 m)above the lake (approximately 392 ft or 119 m elevation).From approximately the same horizon (394 ft or 120 m el-evation), Boekenkamp (1963) collected a Ponticeras (CU41075) that is somewhat more involute than typical P. per-latum. These localities were not examined in this study. Gor-don Baird and Carl Brett have identified a Lodi Limestonehorizon with abundant auloporid corals in the basal 3 m (10ft) of this section at approximately 3 m (10 ft) above lakelevel (Gordon Baird, pers. comm., 2002).

LocalityW, I-11c, south of McKinneys.–From a level inthe Sherburne Siltstone at the side of the railroad beginningapproximately 0.5 mi (0.80) south of McKinneys (Tomp-kins County) at the southeastern end of Cayuga Lake, Kin-dle (1896) collected specimens of Ponticeras perlatum: CU10939b and c from his locality 7-2 and CU 10933a and bfrom “a short distance to the south” at his locality 7-3. Thefaunas at Kindle’s localities 7-2 and 7-3 include the aulo-porid coral Cladochonus and could be the faunal horizon ofH. S. William’s (1884) station 48 locality along the side ofthe railroad “a little north of McKinney’s station.” If so, Kin-dle’s (1896) localities would be 15 m (50 ft) below the topof the Sherburne (and base of Renwick Shale) (Williams,1884: 60) or they could be lower horizons, as much as 30 m(100 ft) below the top of the Sherburne. In the section aboveWillow Point (Loc. I-11), which is approximately 0.5 mi(0.80 km) south of McKinneys, de Witt & Colton (1978:table 1) recorded the top of the Sherburne and base of theRenwick at 505.2 ft (155.0 m) elevation. These localitieswere not examined in this study.


Locality X, Dy-10, Fall Creek, Ithaca.–In the classic sec-tion at Fall Creek (Hall, 1843: 250), on the northeasternside of Ithaca (Tompkins County), beginning at the base ofIthaca Falls, the Renwick Shale and much of the overlyingIthaca Shale and Sandstone are exposed in a spectaculargorge. A 0.30 m (1 ft)-thick siltstone cropping out at thelevel of the pool at the base of the falls marks the top of theSherburne Siltstone and from the Warrenella laevis beds inthe lower Renwick Shale approximately 1 m (3 ft) above,H. S. Williams (1884: locality 1-B) collected a specimen re-ferred to Ponticeras perlatum (CU 42289). No new materialwas found at this level here or in the lower Renwick bedsexposed in the Stewart Avenue access road to NY Rte. 13approximately 0.7 mi (1.1 km) to the north (locality illus-trated by Grasso et al., 1986: fig. 16).

De Witt & Colton (1978) recognized 13 m (42 ft) ofRenwick Shale in the section but the contact with the over-lying Ithaca beds is inacessable in the falls. Just above thecrest of the falls on the southern side of the creek is the siteof the Old University Quarry from which S. G. Williamscollected the specimen (NYSM 3753) referred here toChutoceras nundaium. The locality is Williams’ (1884: sec-tion III, station 3), approximately 72 m (235 ft) above the“Spirifer laevis Zone” at the base of the Renwick Shale. Thelithology is the distinctive shelly “Firestone bed” facies(Williams, 1884: 17). In the creek section the horizon proj-ects to the lower part of the “Quarry Sandstone” interval(Kirchgasser, 1985) that forms the cascade behind the oldelectric power plant (Kirchgasser, 1985: 230-231, fig. 5).

A few meters above the top of the power-plant cascade,a prominent 0.61 m (2 ft)-thick siltstone (“Bluestone Bed”of Kirchgasser, 1985) marks the top of the “Quarry Sand-stone” interval. In the first 0.30 m (1 ft) of dark shales im-mediately below the siltstone bed, Kindle (1896) collectedgoniatites from his locality I-6, one of which is a fine spec-imen of Koenenites styliophilus (CU 10937; G115). Addi-tional specimens referred here to K. styliophilus (WTK3929/1) have been collected from the 0.30 m (1 ft) of darkshales beneath the siltstone. The horizon, 82 m (270 ft)above the base of the Renwick, is believed to be the LindenHorizon of the upper Penn Yan Shale in western sections(Kirchgasser, 1985). Others specimens of K. styliophilus fromhigher in the section are NYSM 12165/WTK 3931/1, aspecimen found in float on top of the siltstone ledge, and aspecimen (CU 10925) collected by Kindle (1896) from hislocality I-7 in the floor of the creek above the siltstone. Ad-ditional but poor specimens of Koenenites include CU10929 from Kindle’s locality I-8, just below the base ofTriphammer Falls. The upper part of section was illustratedby Kirchgasser (1985: fig. 5; see Addendum).

Locality Y, Dy-10a, Baker Quarry.–From the Ithaca beds

in the old Baker Quarry on the Cornell University Campusthat was located just aboveWest Road at approximately 710ft (216 m) elevation, a specimen presumably collected by J.W.Wells (CU 40106 from his locality 42) is probably a Koe-nenites, as is an uncataloged specimen in the CU collected byS. M. Herrick (SMH-1). The Baker Quarry horizon wouldappear to correlate with a level within the Triphammer Fallsin the Fall Creek section.

Locality Z, Dy-10b, Cascadilla Creek, Ithaca.–In Cas-cadilla Creek on the eastern side of Ithaca (TompkinsCounty), a section of Ithaca Shale and Sandstone is exposedin the gorge upstream of the College Avenue bridge. Kindle’s(1896) locality 2-1 at the base of the lowest cascade in thegorge is in dark Renwick Shale. From his locality 2-4, in thelower Ithaca beds, 17 m (55 ft) above his locality 2-1, Kindlecollected a specimen (CU 10932) that might be Chutocerasnundaium. Specimens collected by Kindle (1896) fromhigher in the section (CU 10918; Kindle’s locality 2-10) atthe foot of the falls below the College Avenue bridge(Heustis Street bridge and approximately 70 m (225 ft)above Kindle’s locality 2-1) and NYSM/WTK 3922/1, 2and CU 10923a-b from 18 m (60 ft) higher (Kindle’s local-ity 2-13) are indeterminate.

Locality AA, Dy-10c, South Hill, Ithaca.–Two speci-mens assigned here to Chutoceras nundaium were describedby Hall (1879) from a locality in the Ithaca Shale and Sand-stone at South Hill, Ithaca (Tompkins County). NYSM3749 and 3751 (the lectotype) were from the LowerChemung Group “at the inclined plane of the railroad” at ornear Ithaca, New York. The lithology of the enclosing rockis the shelly Firestone type (Williams, 1884: 17) that yieldedthe University Quarry specimen at the edge of Fall Creek,Ithaca (Loc. Dy-10). The locality could be the same as Kin-dle’s (1896: 24) locality 5-2, “Quarry at Inclined plane,”which is 35 m (115 ft) above the Cayuga Street crossing ofSixmile Creek at the base of South Hill. The abandonedquarry at approximately 500 ft (152 m) elevation is locatedbelow the railroad line and above Stone Quarry Road, justto the west of a small northwestward-flowing creek. No newgoniatites were found at the locality.

Locality BB, I-8a, Enfield Glen.–An uncataloged speci-men in the CU collection from above the top of the lowerfalls in Enfield Glen, approximately 4 mi (6.4 km) south-west of Ithaca (Tompkins County), is assigned here withquestion to Chutoceras nundaium. The specimen collectedby K. E. Caster or G. Q. Williams came from within theIthaca Shale and Sandstone given that the base of the 13+m(44+ft) Renwick Shale at approximately 448 ft (136 m) el-evation in the Cayuga Inlet (Loc. I-8, Lick Brook: de Witt& Colton, 1978: table 1) is at least 50 ft (152 m) belowwhere the section begins near the base of the first falls in the


gorge. No attempt was made here to locate the goniatitehorizon.

Locality CC, Gen 13a, Big Salmon Creek, nearGenoa.–A Tornoceras? was noted in the pyrite capping theFir Tree Limestone of the upper Geneseo Shale in the north-ern of two closely spaced westward-flowing tributaries of BigSalmon Creek, southwest of Genoa (Cayuga County). Sam-ple WTK 3978 was given by Gordon Baird. This is locality32 of Baird et al. (1989: fig. 4F, horizon d) and Kirchgasser(1994).

Locality DD, Gen-14a, Big Salmon Creek, Venice Cen-ter.–A goniatite fragment and protoconch of Ponticeras? wasfound in the pyrite capping the Fir Tree Limestone exposednear the mouth of the southern fork of an unnamed west-ward-flowing tributary of Big Salmon Creek, northeast ofVenice Center (Cayuga County). The sample (WTK 3989)was provided to us by G. Baird from locality 31 of Baird etal. (1989: section 4E, horizon d) and Kirchgasser (1994).

Locality 28. Pleasant Valley, near Sherburne.–The Gene-seo Shale crops out in the creek immediately opposite Pleas-ant Valley Bible Baptist Church, in the creek just south ofPleasant Valley Road, 1.9 mi (3.0 km) south-southeast ofSherburne (Chenango County). From here, J. W. Wellsmade a good collection of tornoceratids and ponticeratids,and one specimen of Pharciceras? sp. (NYSM 12128; see Pl.32, Fig. 14) is known. Gordon Baird (pers. comm., 2002)recently visited this locality and recognized a few feet of theFillmore Glen Member of the Tully Limestone at the base ofthe section (probable source of the Pharciceras? sp.), followedby approximately 3 m (12 ft) of Geneseo Shale, which isabruptly succeeded by massive beds of Sherburne Siltstoneexposed upstream of the church area.

Genundewa LimestoneAlthough relatively thin, the Genundewa Limestone (Styli-ola Limestone of early authors) is the only widespread lime-stone unit above the Tully in New York and the oldesthorizon with a rich and diverse goniatite fauna. Unfortu-nately, because of generally poor preservation due to recrys-tallization and sporadic occurrence, the fauna is stillimperfectly known. At the type locality, at GenundewaPoint on the eastern side of Canandaigua Lake, de Witt &Colton (1959) recognized 3.7 m (12 ft) of GenundewaLimestone consisting of several 25-250 mm (1-10 in)-thicknodular styliolinid layers separated by dark gray shale. West-ward toward Lake Erie, with the thinning and eventual lossof the shale beds, the nodular beds converge to form an ir-regularly bedded limestone (informal lower GenundewaLimestone) overlain by a single well-bedded limestone (in-formal upper Genundewa Limestone often exposed as thecaprock of waterfalls. West of Linden (Loc. 15), in Genesee

County, the nodular beds of lower Genundewa are progres-sively cut out leaving only a single well-bedded upper Ge-nundewa horizon exposed in Erie County.

The nodular beds of the Genundewa mark the entry ofTornoceras uniangulare compressum Clarke, 1898, Koenenitesstyliophilus kilfoylei n. ssp., and Acanthoclymenia genundewa(Clarke, 1898). Also there is K.? fasciculatus (Clarke, 1898),which is based on a single specimen collected at GenundewaPoint, Canandiagua Lake (Loc. 23b). The Manticoceraslamed group (sensu House & Ziegler, 1977), in the form ofM. contractum (Clarke, 1898), is now believed to enter thesuccession above the nodular beds, in the more regularlybedded upper Genundewa. M. sinuosum apprimatumClarke, 1898, which is similar toM. contractum, might alsoprove to enter in the topmost bed(s) of the Genundewa. Theposition in the Genundewa ofM. nodifer (Clarke, 1885) isunknown.

Because of the thinness of the Genundewa interval, ex-panded scales on Text-fig. 8 are used in sections west ofWhite Creek (Loc. 16); these are indicated by black dotsand open circles next to the solid vertical band beside thesections. Surfaces of submarine erosion associated with theGenundewa Limestone in the condensed Genesee sectionsnear the basin margin in Erie County have been docu-mented in detail by Brett & Baird (1982, 1990, 1996, 1997)and Baird & Brett (1986a, b) (Text-fig. 7). The sub-Genun-dewa disconformity with the underlying North Evans Lime-stone (Conodont bed of Hinde, 1879), which is at the baseof the upper Genundewa in Erie County, serves to separatethe nodular lower Genundewa from the more regularly bed-ded upper Genundewa farther east between Bethany Center(Loc. 24), Linden (Loc. 15), and Linden Falls (Loc. 15a) inGenesee County, and Fall Brook (Loc. 19) in the GeneseeValley in Livingston County. Between the Genesee Valleyand the type section at Canandaigua Lake, the shale in-terbeds in the Genundewa interval thicken and the differen-tiation of the Genundewa into lower and upper parts is lessapparent.

East of Canandaigua Lake where the Penn Yan Shale andWest River Shale pass into the slope facies of the Ithaca Shaleand Sandstone, the Genundewa Limestone horizon is diffi-cult to locate. At Shuman Cemetery, approximately midwaybetween Canandaigua Lake and Keuka Lake, the Genun-dewa is represented by a single styliolinid bed several cmthick (locality Ph-5 of de Witt & Colton, 1978: pl. 4) andcapped by a 1 in (25 mm)-thick lag bed. This styliolinid bedwas thought by deWitt & Colton (1978) and earlier work-ers to be the easternmost occurrence of the unit. FromKeuka Lake southeastward to Cayuga Lake, de Witt &Colton (1959, 1978: pls 3-4) correlated the Genundewawith the Crosby Sandstone of Fox (1932) and Torrey et al.


(1932), and used the base of the Crosby as a datum withinthe thick and undifferentiated silty faces of the Ithaca Shaleand Sandstone. As noted earlier (Kirchgasser, 1985), Gor-don Baird discovered the Genundewa horizon at KeukaLake in Ithaca facies 5-9 m (16-30 ft) above the Crosby. AtPenn Yan (DDs; Py-10/1), Willow Grove (EE; Py-14/2),and Sunset Point (FF; Py-16/2), the bed is a pyritic, nodulardiscontinuity horizon with goniatites and conodonts thatare restricted to the lower and upper Genundewa fartherwest.

Still farther southeast at Seneca Lake, in the creek east ofStarkey (Yates County), a 5-cm-thick, cross-laminated andpyritic styliolinid bed identified by Gordon Baird as Genun-dewa lies within Ithaca facies 16 m (53 ft) above the base ofthe Crosby. No goniatites have been found in the bed, whichcrops out in the creek approximately 30 yd (27 m) south ofNY Rte. 14 (locality GG; OV-6 of de Witt & Colton,1978). It is unclear whether the horizons identified as Ge-nundewa at Keuka Lake and Seneca Lake represent a singlecondensed record of the entire Genundewa interval to thewest or only one of possibly several cryptic Genundewa hori-zons that splay into the southeastward-thickening IthacaShale and Sandstone; for discussion of the genesis of thesomewhat similar Fir Tree and Lodi Limestone horizons, seeBaird et al. (1989). The position of the Genundewa and itsfauna is unknown in the Ithaca Shale and Sandstone atCayuga Lake but the interval probably projects to a levelhigh in that unit (Kirchgasser, 1985: fig. 3).

Localities.–Locality numbers with asterisks have sections il-lustrated in Text-figs 8-9.

Locality 13*, Cayuga Creek.–A poorly preserved Koe-nenites was noted in the topmost 25 mm (1 in) shaley layer(Bed 13/9) of the 0.2 m (8 in)-thick Genundewa Limestoneat the top of the falls over the Genundewa in Cayuga Creek,2 mi (3.2 km) northwest of Cowlesville (Wyoming County).

Locality 14a, Murder Creek.–In Murder Creek at Gris-wold (Genesee County), 0.8 mi (1.3 km) south of Darien,goniatites are abundant in the basal bed, 51-76 mm (2-3 in)thick, of the Genundewa Limestone (Bed 14a/3), whichcrops out at the level of a farm road that crosses the stream34 m (37 yd) upstream of the culvert beneath the railroadtracks. This locality was noted by Clarke (1899c: 108-109).

Locality 15*, Linden.–The Genundewa Limestoneforms the caprock of the waterfall in the small eastward-flowing tributary to Little Tonawanda Creek, 0.3 mi (0.48km) north of the bridge at Linden (Genesee County). In the33-cm-thick exposure a few yards (meters) south of the falls,Koenenites and Tornoceras are common in the 10 cm nodularbed at the base of the unit. At the top of the overlying 5 cmnodular bed, pyritic bactritids and protoconchs and earlywhorls of Koenenites occur in microfossil residues from the

presumed North Evans Limestone (Kirchgasser, 2002), alevel marking the hiatus between the 23-cm-thick nodularlower Genundewa and the 18-cm-thick, more regularly bed-ded, upper Genundewa.

Locality 15a, Linden Falls.–At the main falls in the LittleTonawanda Creek at Linden (Genesee County), the Genun-dewa Limestone is 0.36 m (14 in) thick. Goniatites areabundant in Bed 15a/12, the lowermost 76 mm (3 in) ofthe 0.18 m (7 in)-thick nodular lower Genundewa and inthe overlying Bed 15a/13, a 76 mm (3 in)-thick bed in themiddle of the lower Genundewa interval. Only a few goni-atites were noted in the more uniformly bedded and resist-ant 0.18 m (7 in)-thick interval of the upper Genundewathat caps the falls.

Locality 16*, White Creek.–A 0.34 m (1.1 ft)-thick sec-tion of Genundewa Limestone crops out in the bed of thewestern branch of White Creek in a field north of NY Rte.20, 2.1 mi (3.4 km) east of Bethany Center (GeneseeCounty). A few poor specimens of Koenenites were noted inBed 16/8, the 0.08-0.09 m (3-4 in)-thick lower nodular in-terval of the Genundewa. The upper, more regularly beddedinterval is 0.26 m (10 in) thick and poorly preserved bariticgoniatites occur in Bed 16/9, the uppermost 76 mm (3 in)of the unit.

Locality 17*, Taunton Gully.–At the top of the upperfalls in Taunton Gully, 1.7 mi (2.7 km) north of Leicester(Livingston County), below the bridge where Starr Roadcrosses the creek, the Genundewa Limestone is 0.56 m (1 ft10 in) thick. Above a 0.13 m (5 in)-thick black shale thatmarks the top of the Penn Yan Shale, the lowermost 0.25 m(10 in) of the Genundewa consists of nodular limestone.Bed 17/8 at the top of this nodular interval is the level thatproduces Acanthoclymenia genundewa and Tornoceras. Theoverlying 0.25 m (10 in) of more uniformly bedded upperGenundewa is sample-horizon 4586-SD of de Witt &Colton (1978) and Huddle (1981: B53).

Locality 18*, Beards Creek.–The Genundewa Limestonecaps the spectacular falls in Beards Creek, approximately 175yd (158 m), downstream from the Dunkley Road bridge,0.8 mi (1.3 km) northwest of Leicester (Livingston County).The nodules of Bed 18/6a produce Acanthoclymenia genun-dewa, rare Koenenites styliophilus kilfoylei n. ssp., andTornoceras; the bed lies 0.15 m (6 in) above the lip of thefalls, at the top of the 0.15 m (6 in)-thick nodular lower Ge-nundewa Limestone; this is locality 3001 of Kirchgasser(1975: 68). The overlying 0.25 m (10 in)-thick upper Ge-nundewa is more massive and regularly bedded.

Locality 19, Fall Brook-Dewey Hill.–At the top of theNY Rte. 39-20A roadcut at Dewey Hill, 1 mi (1.6 km)south of Geneseo (Livingston County), the lowermost 0.25m (10 in) of nodular Genundewa Limestone is exposed on


the southern side of the road above a 0.13 m (5 in)-thickblack shale overlying a prominent 0.1 m (4 in)-thick, well-bedded argillaceous limestone at the top of the Penn Yan.Poor Koenenites and Acanthoclymenia occur in the nodules ofBed 19/10a at the base of the interval; this horizon is bedDH-1 of Kralick (1991, 1994) and is approximately thelevel of sample-horizon 8962-SD of Huddle (1981: B52).Bed 19/10, at the top of the nodular interval, is a 0.05 m (2in)-thick nodule bed with fragments of Manticoceras con-tractum?, Acanthoclymenia, and Tornoceras. Bed 19/10 corre-sponds to bed DH-2 of Kralick (1991, 1994) andsample-horizon 6681-SD of de Witt & Colton (1978) andHuddle (1981: B52).

In the side creek at Fall Brook, adjacent to Dewey Hill,the complete Genundewa Limestone is approximately 0.71m (2.3 ft) thick. In the accessible section along the footpath100 yd (91 m) north of the main falls, Bed 19/10 with Acan-thoclymenia genundewa is a 0.12 m (5 in)-thick nodular bedat the top of the lower nodular interval of the Genundewain the recess beneath the well-bedded 0.17 m (7 in)-thickupper Genundewa; here the base of Bed 19/10 is 0.32 m (1ft) above the base of the Genundewa, and 0.29 m (1 ft)below the top of the Genundewa Limestone as here defined;this is bed FB-1 of Kralick (1991, 1994).

Locality 19a, Fall Brook, loose blocks.–Loose blocks ofGenundewa Limestone occur at the base of the high falls inFall Brook, 1.5 mi (2.4 km) south of Geneseo (LivingstonCounty). The blocks contain baritic specimens assigned toManticoceras contractum and are believed to have come fromthe topmost beds of the Genundewa.

Locality 22, Barnes Gully.–In Barnes Gully, above FosterPoint on the western side of Canandaigua Lake, 2.5 mi (4.0km) southeast of Cheshire (Ontario County), the Genun-dewa Limestone consists of a sequence of styliolinid lime-stone horizons. Bed 22/7, 1.14 m (3.75 ft) below thetopmost of the styliolinid units and 1.2 m (4 ft) above a 0.3m (1 ft)-thick black shale, has barite and poorly preservedgoniatites. Here the black shale band is taken to mark thebase of the Genundewa because this is the horizon that canbe seen below the Genundewa to the west. There are styli-olinid limestone horizons lower than this, both here and,for example, at Seneca Point Creek. The top of the Genun-dewa is at 946 ft (288.3 m) altitude (de Witt & Colton,1978: table 1).

Locality 23, Seneca Point Creek.–In Seneca Point Creek,above Seneca Point on the western side of Canandaigua Lake(Ontario County), the styliolinid limestones that comprisethe Genundewa Limestone extend through an interval of2.4 m (7.9 ft) and are exposed in the face of the first fallsabove the lake (Text-fig. 9). Bed 23/1 is a 0.23 m (9 in)-thick styliolinid unit with barite, which forms the lip of the

falls, 1.25 m (4.1 ft) above a 0.13 m (5 in)-thick black shalethat marks the top of the Penn Yan Shale, and 1.17 m (3.8ft) below the highest styliolinid bed; this is locality 3006a ofKirchgasser (1975: 68). The fauna of Bed 23/1 includesAcanthoclymenia, Koenenites, and Tornoceras. Tornoceras isparticularly abundant in the correlative of Bed 23/1 inMiller’s Creek, Canandaigua Lake, 0.5 mi (0.8 km) southof Barnes Gully (Bed MC-3 of Kralick, 1991: fig. 2; 1994:1395).

Locality 23b, Genundewa Point, GenundewaLake.–This section is along the eastern side of CanandaiguaLake, north of Genundewa Point (cottage number 900),below East Lake Road, 1 mi (1.6 km) north of Vine Valley(Yates County). Tornoceras sp., Acanthocylmenia genundewa,and Koenenites? fasciculatus occur along with abundant bac-tritids in the styliolinid lenses of Bed 23b/4 cropping outbetween 0.71 m (2.3 ft) and 0.84 m (2.75 ft) above the 0.1m (4 in)-thick black shale that defines the base of the 2.4 m(8 ft)-thick Genundewa interval. Tornoceras is relativelycommon in the fossiliferous and baritic styliolinid ledge(Bed 23b/5) that lies between 1.14 and 1.45 m (3.75 and4.75 ft) above the base of the black shale at the base of theGenundewa; this is sample-horizon 6718-SD of de Witt &Colton (1978: Section Nap-4, pl. 4) and Huddle (1981:B51) and the correlative of Bed 23/1 in the Seneca Pointsection. Poor baritic goniatites were noted in the uppermoststyliolinid ledge (Bed 23b/6), which dips below lake levelapproximately 50 yd (46 m) north of the point. Bed 23b/6(=3958-SD) lies between 1.96 and 2.13 m (6.4 and 7 ft)above the base of the Genundewa; the top of Bed 23b/6 im-mediately underlies a 0.3 m (1 ft)-thick interval of thin styli-olinid lenses that is taken as the top of the GenundewaLimestone.

Locality 24, Bethany Center.–Numerous loose blocks ofGenundewa Limestone cover the embankment below NYRte. 20 for several hundred yards (meters) east of the CenterRoad bridge that crosses NY Rte. 20 at Bethany Center(Genesee County). The goniatites Koenenites styliophilus kil-foylei n. ssp. and Tornoceras uniangulare compressum areabundant in certain levels, but unfortunately most of theblocks are highly weathered and the already recrystallizedspecimens tend to crumble. The most productive horizon isBed 24b/1 of the Genundewa outcrop below the CenterRoad overpass (Loc. 24b). Small pyritic specimens of Acan-thoclymenia occur in the shaley styliolinid beds (Bed 24/2)on the tops of loose blocks on the southern side of the high-way (including specimens donated by James Kradyna). Thehorizon is probably from the topmost 76 mm (3 in) of theGenundewa exposed beneath the bridge.

Locality 24b, Bethany Center roadcut.–The upper bedsof the Penn Yan and the overlying Genundewa Limestone


are seen in the roadcut below the Center Road overpass ofNY Rte. 20 at Bethany Center (Genesee County). The Ge-nundewa exposure is directly beneath the bridge, and herethe limestone is 0.38-0.40 m (1.25 ft) thick. Goniatites aremost abundant in the 0.12-0.15 m (5-6 in)-thick nodularlower Genundewa (Bed 24b/1).

Locality DD, Py-10, Cornwall Gully.–In CornwallGully, above NY Rte. 364 on the northwestern side of PennYan (Yates County), the Genundewa Limestone is repre-sented by a pyritic lag deposit (Bed Py-10/1) within atongue of the Ithaca Shale and Sandstone. The horizon is athin band of rusty pyritic nodules with poorly preservedKoenenites? fasciculatus and Acanthoclymenia within a 0.79m (2.6 ft)-thick shaly interval, which crops out on the east-ern side of the creek above a series of low falls over siltstones,between 5.2 and 6.1 m (17 and 20 ft) above the base of theCrosby Sandstone. The 2 m (7 ft)-thick Crosby Sandstoneat the base of the tongue of the Ithaca Shale and Sandstonecrops out at approximately 860 ft (262 m) elevation (deWitt & Colton, 1978: table 1, pl. 4). This locality was foundby G. Baird.

Locality EE, Py-14, Willow Grove Gully.–In WillowGrove Gully, Keuka Lake, above NY Rte. 54, 3.5 mi (5.6km) southwest of Penn Yan (Yates County), the horizon ofthe Genundewa Limestone is represented by Bed Py-14/2,a thin concretionary bed, with pyritic seams and cross sec-tions of Koenenites, that lies within a tongue of the IthacaShale and Sandstone, approximately 8.8 m (29 ft) above thebase of the Crosby Sandstone. The horizon is 0.22 m (9 in)below a prominent 0.34 m (1.2 ft)-thick black shale; theblack shale is the first black shale above the Crosby Sand-stone illustrated by de Witt & Colton (1978: pl. 4).

Locality FF, Py-16, Sunset Point Gully.–In Sunset PointGully, Keuka Lake, above NY Rte. 54, 5.8 mi (9.3 km)southeast of Penn Yan (Yates County), the horizon of theGenundewa Limestone is represented by a thin concre-tionary bed (Py-16/2) with pyritic seams within a tongue ofthe Ithaca Shale and Sandstone, approximately 8.5 m (28ft) above the base of the Crosby Sandstone. Small pyritic go-niatites are common in the bed and include the typical Ge-nundewa forms Koenenites? fasciculatus, Acanthoclymenia cf.genundewa, and Tornoceras. Bed Py-16/2 is 0.91 m (3 ft)below a 0.48 m (1.6 ft)-thick black shale that is the firstblack shale above the Crosby Sandstone illustrated by deWitt & Colton (1978: pl. 4).

Locality GG, Ov-6, creek east of Starkey.–In the creekeast of Starkey (Yates County), and south of Brewer Point,Seneca Lake, G. Baird located the Genundewa Limestoneequivalent at a position 16 m (53 ft) above the base of the1.4 m (4.6 ft)-thick Crosby Sandstone. The horizon is a 5cm (2 in)-thick cross-laminated, pyritic, styliolinid lime-

stone that crops out in the creek 2 m (7 ft) below a massivesiltstone and approximately 30 yd (27 m) south of NY Rte.14. DeWitt & Colton (1978: table 1, pl. 5) recognized 0.46m (1.5 ft) of Crosby Sandstone at this locality with the basecropping out at 577.5 ft (176.0 m) elevation. No goniatiteswere found in the Genundewa at this locality.

West River ShaleThis unit comprises the sequence of medium gray shales andmudrock with numerous black bands that lies between theGenundewa Limestone and the black Middlesex Shale. Go-niatites occur both as pyritic internal molds and moldswithin the common concretions, approximately 0.3 m (1 ft)in diameter, of which there are several levels in the sequence.Goniatites also commonly occur crushed flat within theshale. The unit is 2.4 m (8 ft) thick on Lake Erie near PikeCreek outlet; on the western side of the Genesee Valley it is16.2 m (53 ft) thick and at Seneca Point Creek approxi-mately 40 m (132 ft) thick. East of Canandaigua Lake, theWest River Shale interfingers with the upper part of theIthaca Shale and Sandstone and partly overlies that unit.The West River Shale is more extensive than as originallydefined by Clarke & Luther (1904); the current definitionis that of de Witt & Colton (1959: 2819). Full details ofthickness and lithological variation were given by de Witt& Colton (1978). Meter-scale cycles of alternating blackand gray shale with concretions characterize the West RiverShale particularly between the Genesee Valley and KeukaLake. On the correlation charts of deWitt & Colton (1978:pls 2, 4), 23 cycles can be recognized from no. 1 above thetop of the Genundewa Limestone to no. 23 immediatelybelow the base of the Middlesex Shale.

A useful lithic marker in the upper West River Memberis the Bluff Point Siltstone Bed (Torrey et al., 1932; Bergin,1964; deWitt & Colton, 1978). This horizon, with distinc-tive convolute bedding, ranges in thickness from 1.5 to 11cm (0.05 ft to 0.35 ft) (de Witt & Colton, 1978: table 1).It lies within the fourteenth cycle and can be seen in mostsections between the Genesee Valley and Seneca Lake.

The following sequence of goniatities is recognized inthe area between the Genesee Valley and Keuka Lake. Koe-nenites sp., Acanthoclymenia sp., and Tornoceras sp. occursporadically through the lower third of the member. Thefauna of the middle part of the member up to and includingthe relatively fossiliferous shales surrounding the concre-tionary bed, in the thirteenth cycle, just below the thin dis-tinctive convolute-bedded Bluff Point Siltstone Bed (Bergin,1964; deWitt & Colton, 1978), is dominated by Koenenitesbeckeri n. sp. along with K.? cf. fasciculatus, and Loboto-rnoceras aff. hassoni House, 1978.

The uppermost West River Member contains a distinct


goniatite fauna, particularly in the septarian horizons suchas those 2.4 and 3.4 m (8 and 11 ft) below the MiddlesexShale in Beards Creek (Loc. 18a/4b in cycle 18, theWilliamsburgh Bed of Over et al., 1999, 2003, and 18a/5 incycle 19–20, the Beards Creek Horizon of Kirchgasser &House, 1981). Here are found K. beckeri n. sp. form D,Manticoceras lamed aff. cordatum (G. & F. Sandberger,1850), Acanthoclymenia aff. neapolitana (Clarke, 1892),Acanthoclymenia sp. A (Kirchgasser, 1975; Loc. 26a/1), Pon-ticeras? sp., K.? cf. fasciculatus, and Tornoceras sp. Bed 18a/4bat Beards Creek, named the Williamburgh Bed by Over etal. (1999: B15; 2003: 219) is a styliolinid grainstone, rich inpyrite, wood, and conodonts, which is intermittently ex-posed near the top of the West River Shale between LakeErie and the Genesee Valley. To the east, goniatites are foundin theWilliamsburgh Bed atWhetstone Brook (Bed 26a/1),near Honeoye (Ontario County); the same horizon might bePy-7/2 in Chidsey Point Creek, Keuka Lake, in SteubenCounty.

Localities.–Locality numbers with asterisks have sections il-lustrated in Text-fig. 9. The goniatites from the West RiverShale are mostly poorly preserved and squashed flat, apartfrom a few horizons with pyritic inner whorls and calcitic orbaritic replacements. The distribution of goniatites in thisunit has not been investigated comprehensively.

Locality 18a*, Beards Creek.–The Beards Creek sectioncontinues upward from that illustrated in Text-fig. 8 (Loc.18) and the levels that yielded goniatites are indicated inText-fig. 9. Beds 18a/1-3 are exposed below the DunkleyRoad bridge. Bed 18a/1 is 0.15 m (6 in) above the Genun-dewa, Bed 18a/2, 0.6 m (2 ft) above, and Bed 18a/3 approx-imately 2.13 m (7 ft) above, and 0.3 m (1 ft) below thefoundation of the bridge on the eastern side of the road.Above the road, Bed 18a/4 is a line of large septaria noduleswith goniatites in adjacent shales beneath a 0.17 m (7 in)-thick black shale approximately 7 m (23 ft) below the Mid-dlesex Shale; the bed is in the cycle below the one thatincludes the Bluff Point Siltstone (deWitt & Colton, 1978:pls 2, 6) but the key marker horizon was not seen in the sec-tion. In the succeeding cycle, which elsewhere includes theBluff Point Siltstone, Bed 18a/4a (cycle 15) is a graymudrock with Koenenites beckeri n. sp., 4.9 m (16 ft) belowthe base of the Middlesex Shale. Near the top of the section,Bed 18a/4b (Williamsburgh Bed) in cycle 18 is a Fossil Logbed (concretionized raft of driftwood), 3.4 m (11 ft) belowthe base of the Middlesex Shale which underlies a 0.3 (1 ft)black shale; this linguloid-rich bed yieldsManticoceras lamedaff. cordatum, K.? cf. fasciculatus, Ponticeras? sp., Acanthocly-menia aff. neapolitana, and Tornoceras. Bed 18a/5, the BeardsCreek Horizon, in cycle 19-20, is a line of vast septaria to

1.5 m (5 ft) across, 2.4 m (8 ft) below the base of the Mid-dlesex Shale. Koenenites beckeri n. sp. form D and Ponticeras?sp. occur in Bed 18a/5.

Locality 19a*, Fall Brook.–The Fall Brook section con-tinues upward from that shown inText-fig. 8 (Loc. 19). Bed19a/1 is the shale unit 0.9 m (3 ft) above the Genundewaimmediately above the falls. Beds 19a/2-4 are above thehighway bridge of NY Rte. 63. Bed 19a/2 is approximately0.5 m (2 ft) upward in the exposure above the bridge, per-haps 2.3 m (7.5 ft) above the Genundewa, and the goniatitesare crushed in a capping seam of a thin Styliolina-rich unit;this is sample-horizon 8122-SD of deWitt & Colton (1978:pl. 2) and Huddle (1981: B52) (Kirchgasser et al. 1994: fig.10). Bed 19a/3 is from dark shale approximately 4.7 m (15.5ft) higher, and Bed 19a/4 is a large septarian level approxi-mately 2.0 m (6.5 ft) higher again.

Locality 23a, Seneca Point Creek.–There is a long sec-tion of the West River Shale in Seneca Point Creek, aboveSeneca Point, on the western side of Canandaigua Lake (On-tario County). At the top there is a major falls over the blackMiddlesex Shale. Bed 23a/3 is a concretionary level at thetop of cycle 13, which is overlain by a 0.18 m (7 in)-thickblack shale at the base of cycle 14. The horizon crops outapproximately 60 yd (54 m) downstream from the foot ofthe falls and just above a cascade where a side creek entersfrom the north. Bed 23a/3 is 0.67 m (2.2 ft) below the 1.5cm (0.6 in) Bluff Point Siltstone in cycle 14 (see de Witt &Colton, 1978: pls 3-4). Bed 23a/3 yields Koenenites beckerin. sp. and Lobotornoceras aff. hassoni. Lower, another sidecreek enters from the north, and Bed 23a/2, a concretionhorizon exposed approximately 30 yd (27 m) below this, isin cycle 13, 2.7 m (8.75 ft) below the Bluff Point Siltstone.Farther downstream, a prominent 0.38 m (1.25 ft)-thickblack band is well exposed in the floor of the creek upstreamof a low falls, and Bed 23a/1 with K. sp. is a concretion bedin the cycle below (cycle 12), approximately 5.3 m (17.5 ftbelow the Bluff Point Siltstone. In this section, the BluffPoint Siltstone is approximately 15.5 m (51 ft) below thebase of the Middlesex Shale (de Witt & Colton, 1978).

Locality 26a*, Whetstone Brook.–The upper part of theWest River Shale is exposed in Whetstone Brook down-stream from the crossing of Huff Road (which passes west toBig Tree Road), 1.6 mi (2.5 km) northwest of Honeoye(Ontario County). The section illustrated here below theMiddlesex Shale commences at approximately 1,035 ft (315m) altitude (Text-fig. 9). Two horizons consist of a large-septarian nodule horizon (Bed 26a/2) above, and a smallernodule horizon (Bed 26a/1), 0.76 (2.5 ft) below a 0.3 m (1ft)-thick black shale unit; Bed 26a/1, with Acanthoclymeniasp. A and Lobotornoceras aff. hassoni, is locality 3003 ofKirchgasser (1975: 68) and theWilliamsburgh Bed of Over



et al. (1999, 2003). The key black shale, at the base of WestRiver cycle 18, lies 6.1 m (20 ft) below the base of the Mid-dlesex Shale (de Witt & Colton, 1978: pl. 2) and can betraced on the correlation charts of deWitt & Colton (1978:pls 2, 4) eastward to Canandaigua Lake and westward to theGenesee Valley where at Beards Creek (Loc. 18a) it is theboundary between cycle 18 with the Fossil Log concre-tionary bed (18a/4b) and the Beards Creek Horizon (18a/5)in cycle 19-20 (Text-fig. 9). Poor pyritic goniatites occur inthe dark shales of Bed 26a/3, approximately 0.91 m (3 ft)below the base of the Middlesex Shale.

Locality 29a, Hicks Point.–This is the locality on thewestern side of Canandaigua Lake (Ontario County) fromwhich Wells (1956b) reported the first specimen of thegenus Koenenites known in New York. The section at thesouthwestern side of Hicks Point begins at lake level andcontinues in the bluffs below and above Bopple Hill Road.The locality was visited by MRH with J. W. Wells in 1959,and, although the specimen was loose, it was thought itcould have come from the shale interval 5.5-7.3 m (18-24ft) above the lake level, and below the two septarian horizonshigher in the cliff. It could also have come from Bed 29a/1,a 0.46 m (1.5 ft)-thick band of dark gray to black silty shaleapproximately 2.6 m (8.5 ft) above lake level that yields sim-ilarly preserved crushed molds of Koenenites and Acanthocly-menia. Bed 29a/1 is within the section of gray mudrockexposed in the shale pit below the road and just west of thecreek.

Locality 46c, Snyder’s Gully.–The upper beds of theWest River Shale are accessible in Snyder’s or Snyder Gullyat the southern end of Canandaigua Lake, above NY Rte.21, 0.3 mi (0.5 km) south of Woodville (Ontario County).Bed 46c/1, with Koenenites beckeri n. sp., is a 0.22 m (8.5in)-thick Fossil Log concretionary bed 5 m (16.4 ft) belowthe base of the Middlesex Black Shale. The horizon is 0.9 m(3 ft) below a 0.1 m (4 in)-thick black shale and approxi-mately 4 m (13 ft) above where continuous exposure begins.

Locality KK, Py-16, Sunset Point Gully.–In the upperpart of the section in Sunset Point Gully, Keuka Lake, poorlypreserved Koenenites occur in Bed Py-16/3, a horizon ofnodules 0.91 m (3 ft) below the Bluff Point Siltstone, andpoorly preserved Koenenites, Acanthoclymenia, and Tornocerasoccur in the soft shales surrounding a 0.15 m (6 in)-thickconcretionary horizon (Py-16/4) beneath a 0.1 m (4 in)-thick black shale 5.4 m (17.6 ft) above the Bluff Point Silt-stone. The Bluff Point Siltstone is well exposed in the section(de Witt & Colton, 1978: table 1, pl. 4).

Locality LL, OV-10, Mill Creek.–In the upper part ofMill Creek near Lodi (Seneca County), approximately 145yd (133 m) downstream of the NY Rte. 414 bridge, the go-niatite bed with Koenenites beckeri n. sp. in the cycle below

the Bluff Point Siltstone (Bed Ov-10/6) is a dark gray shaleimmediately above a 10 in (0.25 m)-thick black shale. Thehorizon, at approximately 905 ft (276 m) elevation, is 3.6 m(11.8 ft) below the Bluff Point Siltstone. The black shaleand overlying Bluff Point Siltstone were illustrated by deWitt & Colton (1978: pl. 3); see also Oliver & Klapper(1981b: 45, fig. 1, sample WTK 3937).

Locality HH, Nap-3, Granger Point Road.–The upperWest River Shale is well exposed in the roadcut and sidecreek along NY Rte. 21 across from the road to GrangerPoint, Canandaigua Lake (Ontario County). The BluffPoint Siltstone is approximately 4 m (13 ft) above a 0.5 m(1.6 ft)-thick dark gray to black shale near the base of thesection. The Bluff Point Bed is 0.5 m (1.6 ft) above a 0.15m (6 in)-thick black shale, and specimens of Koenenites indark shales and concretions in the float at the base of thesection might have come from the cycle below the 0.15 m(6 in)-thick black shale.

Locality II(XX), Nap-8, 0.8 mi south ofMiddlesex.–This section in the West River Shale is in thewestward-flowing creek that crosses NY Rte. 245, 0.8 mi(1.3 km) south of Middlesex (Yates County). Approximately46 ft (14 m) upstream from where the section commencesis a band of small concretions in the creek floor and the 50mm (2 in)-thick convolute-bedded Bluff Point Siltstone lies6.3 m (21 ft) above. Bed Nap-8/1 (WTK 3884) is a fossil-log type concretion with pyrite and barite seams with afauna of styliolinids, gastropods, small lingulid brachiopods,and goniatites. The loose concretion is believed to havecome from the line of concretions 3 m (9.8 ft) below theBluff Point Siltstone; the horizon lies between two 80 mm(3 in)-thick black shales that define the first cycle below thecycle with the Bluff Point bed. Goniatites in Nap-8/1 in-clude Koenenities beckeri n. sp., K.? cf. fasciculatus, Tornocerassp., and ?Lobotornoceras aff. hassoni. Nap-8/2 is a graymudrock with goniatite molds, 0.61 m (2 ft) above Nap-8/1 and just below the upper 80 mm (3 in)-thick blackshale.

Locality JJ, Py-7, Chidsey Point.–There is an extensivesection of the West River Shale in Chidsey Point Creek onthe West Branch of Keuka Lake in Steuben County, 1.7 mi(2.7 km) south of Branchport (Yates County). Bed Py-7/1is a nodular silty mudstone with crushed molds approxi-mately 4.6 m (15 ft) below the base of the Bluff Point Silt-stone; this is sample-horizon 8556-SD of deWitt & Colton(1978: pl. 6) and Huddle (1981: B50). No goniatites wererecovered from the cycle below the one with the Bluff PointSiltstone. Bed Py-7/2 is a Fossil Log concretionary bed withsmall pyritic and baritic goniatites including Koenenites beck-eri n. sp. form D, Acanthoclymenia aff. neapolitana, andTornoceras, 3.4 m (11 ft) below the base of the Middlesex

Shale exposed near the top of the section where the creeknarrows near the top of a 3 m (10 ft)-high cascade. The hori-zon is 1.5 m (5 ft) above a series of three thin siltstones. BedPy-7/2 might correlate westward with the Fossil Log con-cretionary bed 18a/4b, the Williamsburgh Bed of Over etal. (1999, 2003), in Beards Creek (Loc. 18a) in the GeneseeValley.

Locality MM, Gen 1a, Sheldrake Creek.–In SheldrakeCreek, on the western side of Cayuga Lake, Bed Gen 1a/1is the goniatite bed with Koenenites beckeri n. sp. in the cyclebelow the Bluff Point Siltstone. The horizon is a gray shaleimmediately below a 0.91 m (3 ft)-thick black shale crop-

ping out in the creek floor approximately 100 yd (91 m) up-stream of the Munson Road bridge. The 8 cm (3 in)-thickBluff Point Siltstone, which caps a waterfall at a ford cross-ing, lies 2.4 m (8 ft) above the goniatite bed. In the sectionof de Witt & Colton (1978: pl. 3), the cycle that includesthe Bluff Point Siltstone is in the center of the eastward-thinning wedge of West River Shale; the black shale aboveBed Gen 1a/1 is the lower of the two black shales shown intheir section.

Locality NN, I-3, Coy Glen.–In Coy Glen, 2.1 km (1.3mi) west of Ithaca (Tompkins County), Bed I-3/1 is amudrock with a “Naples” fauna of goniatites (Koenenites

Text-fig. 9. Sections of the Penn Yan Shale, Genundewa Limestone, andWest River Shale. Section of the Penn Yan Shale and the lower Ge-nundewa Limestone at Seneca Point Creek, Canandaigua Lake, and sections of theWest River Shale between Leicester (Livingston County)and Whetstone Brook (Ontario County), and the locality numbers used in the text.


beckeri n. sp.? and Tornoceras), bactritids, ostracods, andstyliolinids, which could correspond to the goniatite bed inthe cycle below the Bluff Point Siltstone to the west. Thehorizon, at approximately 965 ft (294 m) elevation, is nearthe top of the Ithaca Shale and Sandstone (de Witt &Colton, 1978: pl. 3). It is at the base of a 0.15 m (6 in)-thick dark gray mudrock above a 0.10 m (4 in)-thick blackshale near the top of a low falls, 2.7 m (9 ft) below the levelof the first driveway bridge that crosses the creek upstreamof the Elm Street Extension Bridge. Bed I-3/1 is close to thelevel of sample-horizon 6791SD of de Witt & Colton(1978: pl. 3) and Huddle (1981: B47), which lies down-stream of the Elm Street Extension Bridge.

Locality OO, I-8, Lick Brook.–In Lick Brook, 2.7 mi(4.3 km) south of Ithaca, Bed I-8/1 is a silty mudrock withgoniatites (Koenenites beckeri n. sp.) in the Ithaca Shale andSandstone that could correspond to Bed I-3/1 in Coy Glen.Bed I-8/1, at approximately 980 ft (298 m) elevation, is atthe base of a 0.1 m (4 in)-thick silty mudrock overlying a0.05 m (2 in)-thick siltstone and a 0.1 m (4 in)-thick blackshale that forms a recess in the section on the western sideof the creek approximately 100 yd (91 m) downstream ofthe pipeline crossing. The Sand Bank Road crossing is far-ther downstream. The black shale could be the one illus-trated by de Witt & Colton (1978: pl. 3), 47 m (155 ft)below the base of the Middlesex Shale.

SONYEA GROUP

In its type area in the Genesee Valley and westward, theSonyea Group (Chadwick, 1933; Rickard, 1975) comprisesthe black Middlesex Shale and overlying green CashaquaShale and is perhaps the most clearly defined of the majorcycles in the New York succession. West of CanandaiguaLake, the uninterrupted green shale facies of the Cashaquacontrasts sharply with the interbedded dark gray and blackshales of the Genesee Group and the microcyclothems ofblack shale, gray shale, and turbidite-siltstones in the suc-ceeding West Falls Group. East of the Genesee Valley, theCashaqua Shale interfingers with and comes to overlie thewestward-thinning wedge of turbidites comprising the Pul-teney Shale (Colton & de Witt, 1958) and Rock StreamSiltstone (Bradley & Pepper, 1938). The Cashaqua Shale isoverlain by the Rhinestreet Shale, the basal black shale ofthe West Falls Group. Sutton et al. (1970), Bowen et al.(1974), and MacDonald (1975) have reported on the shelfand near-shore facies and communities of the SonyeaGroup.

Middlesex ShaleThroughout western New York, the black Middlesex Shale(“lower black band” of older reports) marks the boundary

between the Genesee Group and succeeding strata, whichin early stratigraphic classifications comprised the Portageand, farther east, Naples groups. The unit was formallynamed by Clarke (1903) for exposures near Middlesex(Yates County), and later Chadwick (1933) introduced thename Sonyea for the combined Middlesex and Cashaqua.DeWitt & Colton (1959) recognized approximately 1.8 m(6 ft) of Middlesex Shale at Lake Erie, 4.9 m (16 ft) in theGenesee Valley, 11.3 m (37 ft) at Canandaigua Lake, and amaximum of 22.9 m (75 ft) at Keuka Lake, east of which theunit thins rapidly toward Cayuga Lake. In the Seneca Lakearea and eastward, Sutton et al. (1970) recognized twotongues of black Middlesex Shale separated by an eastward-thickening wedge of silty facies, a lower Montour Shale, andan upper Sawmill Creek Shale. The black shales of the Mid-dlesex were not searched systematically for goniatites duringour survey. The only published record is the specimen ofSandbergeroceras syngonumClarke, 1897, reported by Clarke(1898) from Snyder’s Gully, above Canandaigua Lake nearWoodville (Ontario County; Loc. 46c). H. S. Williams col-lected a specimen (NYSM 12026) referable to S. syngonumfrom probable equivalents of the upper Middlesex Shale inTioga County near Harford Mills (Loc. 64), and we foundloose fragments of the species (NYSM 16577, 16578) atthat locality.

Cashaqua ShaleJames Hall (1840) proposed the name Cashaqua Shale forthe distinctive green shales above the “Genesee Slate” ex-posed along Cashaqua Creek (now Keshequa Creek) aboveSonyea (Livingston County). This is the type locality ofManticoceras sinuosum (Hall, 1843), a form that marks theestablishment in New York of Devonian goniatites compa-rable to the assemblages of Europe. It is in the CashaquaShale that the molluscan Naples Fauna (Clarke, 1898, 1904)achieves its richest development, especially in the region ofNaples (Ontario County) at the head of Canandaigua Lake.The unit thickens eastward from approximately 12.8 m (42ft) at Lake Erie to approximately 46 m (150 ft) in the Gene-see Valley and then thins eastward to 15.2 m (50 ft) nearCanandaigua Lake. Farther southeast, between there andSeneca Lake, the Cashaqua is locally condensed and rangesin thickness from 1.2 to 13.7 m (4 to 45 ft) (Colton & deWitt, 1958; Kirchgasser, 1965) (Text fig. 10).

Following the lithostratigraphic studies of Colton & deWitt (1958) and Sutton (1960), Kirchgasser (1965, et seq.)investigated the Cashaqua faunas, particularly the goniatitesand conodonts, in sections between Lake Erie and KeukaLake (Text-fig. 10). Several thin black shales (designated Athrough F) in the lower and upper parts of the unit and afew distinctive carbonate horizons provide useful markers


for locating goniatite horizons. In sections west of the Gene-see Valley, dark gray shales with septarian nodule horizonsand black shales A and B overlie the black shales of the Mid-dlesex. This lower dark shale facies of the Cashaqua thickenseastwardly and in the Genesee Valley and beyond, interfin-gers with the Pulteney/Rock Stream turbidite wedge. In east-ern sections, the stratigraphic positions of marker black shaleB and the Parrish Limestone above the top beds of the RockStream Siltstone indicate that all but the lowermost beds ofthe Cashaqua were deposited after major Sonyea turbiditedeposition had ceased. Goniatites are rare in the Cashaquabelow marker black shale B, but an important baritizedfauna withManticoceras sinuosum clausium n. ssp. occurs atRandall Gully (Bed 44/3). This earliest Cashaqua goniatitefauna reappears, again baritized, in Whestone Brook (Bed42/5), 6.7 m (22 ft) above the marker black shale.

In western sections, the lower dark shale interval of theCashaqua is succeeded by a monotonous sequence of softolive green shales with numerous concretionary horizons.Between Lake Erie (Loc. 30) and Murder Creek (Loc. 34),Probeloceras lutheri (Clarke, 1885) is particularly abundantin the concretion horizons in the middle of the series. Herealso Aulatornoceras eifliense (Steininger, 1849) occurs locally.The concretionary facies of the Cashaqua extends eastwardinto the thick Cashaqua section in the Genesee Valley buthere goniatites are uncommon and less well preserved.Southeast of Honeoye Lake, the concretionary facies passesinto a condensed sequence of soft nodular green shales andsilty shales (Rye Point Shale of Sutton, 1960) and in thispresumably shallower water facies the Naples Fauna attainsits greatest abundance and diversity, particularly among thebivalves Lunulicardium and Ontaria. In this area goniatitefaunas dominated byManticoceras sinuosum sinuosum are lo-cally common in the nodule beds and thin current-reworkedlimestones, especially the nodular red and green ParrishLimestone (Clarke & Luther, 1904) near Naples. As notedby Clarke (1885), the nodular facies of the Cashaqua is re-markably similar to the Kramenzelkalk facies of the Euro-pean Upper Devonian, a condensed facies of cephalopodlimestone that formed on tectonic submarine highs orSchwellen (Rabien, 1956; Tucker, 1973; Krebs, 1979).Kirchgasser (1965, et seq.) has argued that the nodular faciesof the Cashaqua accumulated over Schwellen developed onthe Rock Stream Siltstone. In New York, this distinctivenodular (Knollenkalk) facies reappears in similar strati-graphic settings in theWest Falls Group, again where wedgesof turbidite siltstones at the slope margin penetrate seawardinto the gray shale facies of the basin (Text-fig. 3).

Across the outcrop belt, the upper 3.0 m (10 ft) of theCashaqua consists of a series of dark gray shales, with thebivalve Pterochaenia locally abundant, interbedded with thin

black shales (D through F) (Upper Cashaqua of Kirchgasser,1975). Goniatites are rare in this interval but horizons withProchorites alveolatus (Glenister, 1958) occur at the base orjust below at Murder Creek (Bed 34/19), Little TonawandaCreek (Beds 35/17 and 35/18), Beards Creek (Bed 38/10),and Hicks Point (Bed 45/17). Near the top of this dark shalefacies and just below the base of the Rhinestreet Shale is theShurtleff Septarian Horizon (Kirchgasser, 1975), a line ofdark styliolinid septaria well known for its rich baritizedfauna in the area in the vinicity of Conesus and Honeoyelakes. At this level, Manticoceras sinuosum sinuosum andTornoceras uniangulare obesum Clarke, 1897, are the mostabundant goniatites in a fauna that includes Acanthocly-menia neapolitana, Prochorites alveolatus, and Aulatornoceraseifliense. The upper Cashaqua marks the recurrence of thedark shale facies with Pterochaenia seen earlier in the PennYan and especiallyWest River Shales of the Genesee Group.The counterpart of the Shurtleff Septarian Horizon in theGenesee Group is the Beards Creek Horizon, a line of sep-tarian nodules (Bed 18a/5) near the top of the West RiverShale, best seen in Beards Creek, 2.4 m (8 ft) below the baseof the Middlesex (Text-fig. 9).

Localities.–Locality numbers with asterisks have sections il-lustrated in Text-fig. 10.

Locality 30*, Eighteenmile Creek.–A complete sectionof Cashaqua Shale, one typical of the area west of the Gene-see Valley, is exposed along Eighteenmile Creek below thevillage of North Evans (Erie County), between the railroadoverpass and the NY Rte. 20 bridge. The lower dark shale fa-cies is accessible in the southern bank where exposures beginapproximately 200 yd (183 m) upstream of the railroadcrossing, and there a few goniatites were recovered fromBeds 30/3 and 30/4. Marker black shale B, which caps a lineof concretions (Bed 30/5) at the top of the dark shale inter-val, 2.2 m (7.25 ft) above the Middlesex Shale, extends east-ward to the Genesee Valley and possibly as far east as RandallGully (Loc. 44) in Ontario County. The lower Cashaquaand contact with the underlying Middlesex Shale were illus-trated by Over et al. (2003: fig. 1).

The soft olive-green shales of the middle Cashaqua areaccessible in the cliffs along the northern bank of the creekopposite North Evans. Within these shales are prominenthorizons of argillaceous concretions (Beds 30/7-9) with fre-quent but generally poorly preserved goniatites (mostly Pro-beloceras). Small specimens of Manticoceras sinuosumclausium n. ssp.? are common in thin lenses within Bed30/10, a concretion horizon at creek level at the base of thesection in the steep south bank downstream of the NY Rte.20 bridge. The carbonate horizons above Bed 30/10 are gen-erally unfossiliferous. The dark shale interval of the upper


Text-fig. 10. Sections of the Cashaqua Shale of the Sonyea Group between Eighteenmile Creek, Lake Erie, and Wagener Glen, on Keuka Lake,Steuben County, with the locality numbers used in the text.



Cashaqua is here approximately 2.7 m (9 ft) thick and in-cludes three thin marker black shales. The lowest of these(D) extends eastward to Little Tonawanda Creek (Loc. 35)at Linden (Genesee County), and the next highest (E) as fareast as Briggs Gully (Loc. 43), Honeoye Lake (OntarioCounty). At 0.9 m (3 ft) above marker bed E and 0.9 m (3ft) below the base of the Rhinestreet Shale is the ShurtleffSeptarian Horizon (Kirchgasser, 1975), a horizon that canbe traced approximately 81 mi (130 km) eastward to Con-klin Gully (Loc. 47), near Naples (Ontario County).

Locality 30a, Pike Creek.–Approximately 6.1 m (20 ft)of the upper Cashaqua Shale is exposed in the bed of PikeCreek north of Lake Shore Road, 1.3 mi (2.1 km) northeastof Jerusalem Corners (Erie County). Fragments of goniatiteswere collected at several levels, notably in Beds 30a/1 and30a/2 (the first concretions above where exposures begin),Bed 30a/5 (a thin limestone 0.9 m or 3 ft immediatelybelow marker black shale D), and the shale horizons (Beds30a/8, 9) immediately below and above marker black shaleE, 1.5 m (5 ft) below the base of the Rhinestreet Shale,which caps a low falls.

Locality 31, Smoke Creek.–Along the South Branch ofSmoke Creek below California Road, 1.6 mi (2.6 km)northwest of Orchard Park (Erie County), a complete sec-tion of Cashaqua Shale crops out 0.5 mi (0.8 km) upstreamof the NY Rte. 20 bridge. The sequence is similar to thatseen at Eighteenmile Creek (Loc. 30). Goniatites were notnoted in the septaria and dark shales of the lower Cashaquaexposed near stream level but Probeloceras lutheri is commonin the series of prominent concretionary horizons and sur-rounding green shales exposed in the steep embankment(Beds 31/4 through 31/7). These units correlate with Beds30/7 to 30/10 at Eighteenmile Creek. The lowest of these(Bed 31/4) is 2.6 m (8.5 ft) above marker black shale B. Theconcretions of Bed 31/5 yield P. lutheri and Aulatornoceraseifliense.

Near the top of the embankment and in the southwest-erly flowing side creek, poorly preserved goniatites were re-covered from two horizons in the dark shale facies. Bed 31/2lies immediately above marker black shale D and Bed 31/14is 0.3 m (1 ft) below the base of the Rhinestreet Shale.

Locality 32*, Cazenovia Creek.–Approximately 15 m(50 ft) of Cashaqua Shale are exposed in the ravine of anorthwesterly flowing tributary to Cazenovia Creek, be-tween the main stream and the intersection of North Davisand Conley Roads, 0.8 mi (1.3 km) south of Spring Brook(Erie County). Goniatites were noted in Bed 32/4, a concre-tion horizon 1.5 m (5 ft) above the 75 mm (3 in)-thickmarker black shale B, and in Bed 32/5, 1.5 m (5 ft) higher,but the best collections (withManticoceras sinuosum, Probe-loceras lutheri, and Aulatornoceras eifliense) come from a series

of concretionary horizons (Beds 32/6–7 and especially32/8–9) in approximately the middle of the section. Thesebeds crop out in the bed of the creek downstream of thepoint where the stream bends abruptly to the west belowBed 32/10, a thin but prominent limestone that forms apavement in the creek floor. This same interval of goniatite-rich concretions is seen in all sections between Lake Erie(Loc. 30) andMurder Creek (Loc. 34). A single specimen ofProchorites alveolatus was recovered from Bed 32/16, a hori-zon of large concretions 0.3 m (1 ft) below marker blackshale D at the base of the dark shale interval of theCashaqua. The lower beds of the Rhinestreet Shale are ex-posed in the face of the falls below the road culvert.

Locality 33*, Cayuga Creek.–A long section through themiddle and upper Cashaqua Shale is exposed in CayugaCreek west of Cowlesville (Wyoming County), with expo-sures beginning at approximately 876 ft (267 m) elevation.Goniatites are especially common in a series of concre-tionary levels (Beds 33/4 to 33/7) through a 1.2 m (4 ft)-thick interval of shale that crops out in the floor of thestream near the northern bank, approximately 430 yd (0.4km) downstream of the point where the Erie/Wyomingcounty line crosses the creek and below where NY Rte. 354turns abruptly from southeast to east. The most productivehorizon is Bed 33/5 in which pyritic goniatites (especiallyProbeloceras lutheri) are associated with plant fragments (fos-sil logs).

Higher and less fossiliferous units of the Cashaqua andthe contact with the Rhinestreet Shale are well exposed up-stream along the southern bank near the county line. ThereBed 33/8 is the thin concretionary limestone exposed nearthe water level and Bed 33/9 is the prominent limestone atthe base of the embankment. Bed 33/15 is the prominentledge of limestone 1.7 m (5.5 ft) below marker black shaleD. Bed 33/12 outcrops at stream level where a southward-flowing tributary enters from the north at approximately902 ft (274 m) elevation.

Locality 34*, Murder Creek.–Several goniatite horizonscrop out in the 24.7 m (81 ft)-thick section of CashaquaShale in the ravine of Murder Creek, above Griswold, 1.5 mi(2.4 km) south of Darien City (Genesee County). TheCashaqua/Rhinestreet contact forms the crest of the fallswest of Chick Road at approximately 1,122 ft (342 m) ele-vation. The lowermost 6.1 m (20 ft) of Cashaqua Shale isonly intermittently exposed. Goniatite records begin abovethis interval, in the concretions of Bed 34/3, which underliesmarker black shale B. Bed 34/5 withManticoceras sinuosumsinuosum is a horizon of enormous concretions, to 1.1 m(3.5 ft) long, 1.7 m (5.5 ft) above the black shale; Bed 34/6,with Aulatornoceras eifliense is a smilar horizon with lenses ofmicrofossils, which is well exposed in the creek floor, 1.7 m


(5.5 ft) above Bed 34/5. Upstream is Bed 34/7, the mostproductive horizon (withM. sinuosum sinuosm and Probelo-ceras lutheri) in the section, a layer of disk-shaped concre-tions with unusual, knobby upper surfaces, which crops outat the base of the western embankment in which Beds 34/8through 34/10 are prominently exposed.

Bed 34/12 is a horizon of variably sized concretions withrich microfossil lenses and common goniatites (Manticocerassinuosum sinuosum and Probeloceras lutheri), which crops outin the bed of the stream just below the point where a west-erly flowing side creek joins the main stream below the rightangle bend in Chick Road (1,168 ft or 356.0 m elevationmark). Bed 34/13 is a 76 mm (3 in)-thick limestone-concre-tion horizon at the confluence. Near the top of the green-shale interval of the Cashaqua, goniatites are present butuncommon in Bed 34/17, a thin limestone with Aula-tornoceras outcropping at the base of the falls. Above, Bed34/19 is a layer of large concretions with occasional goni-atites (Prochorites alveolatus) 0.69 m (2.25 ft) below markerblack shale D. A few goniatites (?Probeloceras lutheri) werealso recovered from the dark shales of Bed 34/21 beneathmarker black shale E.

Locality 35*, Little Tonawanda Creek.–A complete sec-tion of Cashaqua Shale crops out in a tributary to LittleTonawanda Creek below the railroad tracks approximately1.3 mi (2.1 km) northwest of Linden (Genesee County).Goniatite records (Probeloceras lutheri) begin in approxi-mately the middle of the section in Bed 35/5, a thin butprominent limestone-concretion horizon 2.4 m (8 ft) belowthe entry of a side creek from the southeast. More produc-tive goniatite horizons are in the concretions of Bed 35/7 atthe entry of the side creek and especially Bed 35/8, withManticoceras sinuosum and P. lutheri, 0.3 m (1 ft) higher.Upstream, the prominent limestone-concretion layer at thebase of the main western embankment is Bed 35/9. In theupper Cashaqua, pyritic specimens of Prochorites alveolatusoccur in the dark shales (Beds 35/17-18) just below andabove marker black shale D.

Locality 37, Wyoming Gulf.–The middle and upperCashaqua Shale are exposed inWyoming Gulf along VillageBrook and in roadcuts on the northeastern side of EastBethany Road, above the village of Wyoming (WyomingCounty). The Cashaqua/Rhinestreet contact is exposed inthe roadcut above the intersection with Wass Road. Goni-atites are scattered through the approximately 24.4 m (80 ft)of section but two horizons produce the best faunas. Bed37/5 is a distinctive horizon of very large concretions (to 0.6m or 2 ft long) with abundant wood fragments, lenses ofmicrofossils, pyrite, and occasionally well-preserved goni-atites, includingManticoceras sinuosum and Tornoceras uni-angulare obesum. The horizon crops out in the bed of the

creek approximately 3.7 m (12 ft) above where exposuresbegin and 4.9 m (16 ft) below the top of a concrete retainingwall bordering the southern side of the road at the edge ofthe stream. A thin black shale noted by Colton & de Witt(1958) approximately 0.9 m (3 ft) below the goniatite hori-zon was not seen. Goniatites (Probeloceras lutheri) also occurin the concretions of Bed 37/7 (Loc. 3008-7 of Kirchgasser,1975) and in Bed 37/8 (withM. sinuosum sinuosum and P.lutheri), 1.7 m (5.5 ft) and 1 m (3.25 ft) below the top of thewall.

Across the road from the concrete wall is a succession ofgoniatite horizons (Beds 37/9 through 37/15) through a 3m (10 ft)-thick interval in the lower part of the lower of tworoadcuts along East Bethany Road. This interval of relativelyabundant goniatites is the easternmost expression of themid-Cashaqua goniatite “zone” noted in sections fartherwest. Bed 37/10 is the prominent limestone near the bottomof the embankment and Probeloceras lutheri is especiallyabundant in the underlying meter of shales (Bed 37/9). Beds37/11-12, with Manticoceras sinuosum sinuosum and P.lutheri, are a pair of concretionary horizons 0.6 m (2 ft)above Bed 37/10; Bed 37/13 is a concretion horizon with P.lutheri and Tornoceras uniangulare in the surrounding shales,0.76 m (2.5 ft) higher, and Bed 37/14 is a shale horizon withP. lutheri and Aulatornoceras, 0.9 m (3 ft) above. Bed 37/15is a concretionary level, with T. uniangulare obesum, 0.61 m(2 ft) above Bed 37/14.

Locality 38*, Beards Creek.–The upper 12.2 m (40 ft)Cashaqua Shale is exposed in the main channel of BeardsCreek, 0.64 km (0.4 mi) northwest of Pine Tavern Corners(Livingston County). A diverse, well-preserved goniatitefauna was collected from pyritic green shales (Bed 38/2) sur-rounding a discontinuous horizon of concretion and nod-ules in the low southern bank of the creek, 1.7 m (5.5 ft)above where continous exposure begins and 10.5 m (34.5 ft)below the Cashaqua/Rhinestreet contact. Bed 38/2 yieldsManticoceras sinuosum sinuosum, Probeloceras lutheri, andAulatornoceras. In the upper Cashaqua, Acanthoclymenianeapolitana was recovered from dark gray shales (Bed 38/10)beneath marker black shale E, and a few goniatite fragmentswere noted in the Shurtleff Septarian Horizon (Bed 38/11).

Locality 39*, Buck Run Creek.–The Cashaqua Shale at-tains its greatest thickness, approximately 45.7 m (150 ft),in the Genesee Valley. The best section for collecting is alongBuck Run Creek, southwest of Mount Morris (LivingstonCounty), beginning several hundred yards (meters) west ofNY Rte. 36. In the lower part of the succession, a 6.4 mm(0.25 in)-thick black shale, 5.8 m (19 ft) above the top ofthe Middlesex Shale and 0.6 m (2 ft) below a horizon of un-usual, small, oddly shaped concretions, is marker black shaleA. Four meters (13 ft) above this black shale is a layer of


small, very dark gray, egg-shaped concretions (Bed 39/3)and goniatites (Manticoceras sinuosum) are first seen in theconcretions of Bed 39/4, which lies 0.6 m (2 ft) above.Marker black shale C is a 2.5 cm (1 in)-thick, prominenthorizon in approximately the middle of the section in thesouthern embankment in the vicinity of a normal fault. Beds39/11 and 39/13 are concretion horizons withM. sinuosumand Aulatornoceras, 1.5 m (5 ft) and 0.61 m (2 ft), respec-tively, below this black shale. Beginning 4.6 m (15 ft) abovethe black shale and continuing for 6.1 m (20 ft) is a succes-sion of concretionary horizons (Bed 39/16 through 39/20)with occasional goniatites, with the best collections comingfrom pyritic shales surrounding Bed 39/17 (with Probeloceraslutheri and Aulatornoceras eifliense ) and Bed 39/19 (withM.sinuosum sinuosum).

The valley narrows near the top of the section and therethe dark shale facies of the upper Cashaqua is exposed in theface of a falls, and above that, the Cashaqua/Rhinestreetcontact is at the top of a narrow flume. Acanthoclymenianeapolitana is found in the 0.6 m (2 ft) of dark shales (Bed39/22), 0.3 m (1 ft) below marker black shale E, and above,baritized specimens (includingManticoceras sinuosum sinu-osum) were recovered from the Shurtleff Septarian Horizon(Bed 39/24), 1.7 m (5.5 ft) above the black shale and 1.4 m(4.5 ft) below the base of the Rhinestreet Shale.

Locality 40*, Cottonwood Point Gully.–Approximately30 m (100 ft) of Cashaqua Shale is exposed in the mainravine west of NY Rte. 256 above Cottonwood Point (Liv-ingston County), on the western side of Conesus Lake. Bed40/1 is a horizon of large concretions in the northern bankat the edge of an open field, just above where exposuresbegin. HereManticoceras sinuosum, Probeloceras lutheri, andTornoceras uniangulare obesum are found in the concretionsthat envelop fossil logs. Goniatites are sporadically distrib-uted through succeeding horizons but near the top of thesection, baritized material (with M. sinuosum sinuosum,Acanthoclymenia neapolitana, and T. uniangulare obesum) isconcentrated in Bed 40/13, the Shurtleff Septarian Horizon,exposed in the face of the falls, 2.1 m (7 ft) above markerblack shale E and 1.1 m (3.75 ft) below the base of theRhinestreet Shale.

Locality 41, Shurtleff ’s Gully.–A section through the up-permost Cashaqua Shale crops out above Cleary Road inShurtleff ’s Gully (Shurger’s Glen), a tributary to KinneyCreek, 1.5 mi (2.4 km) southeast of Livonia (LivingstonCounty). This is the type locality of the Shurtleff SeptarianHorizon (Bed 41/1), which in this area contains the mar-vellously preserved baritized assemblage originally describedby Clarke (1898, 1899a, b). Included are Manticoceras sin-uosum sinuosum, Prochorites alveolatus, Acthoclymenianeapolitana, Aulatornoceras eifliense, and Tornoceras unian-

gulare obesum. The Cashaqua/Rhinestreet contact is at ap-proximately 1,200 ft (366 m) elevation and the septarianbed lies 1.8 m (6 ft) below.

Locality 41a, North McMillan Creek.–The upper 6.1 m(20 ft) of Cashaqua Shale is exposed above East Lake Road,in North McMillan Creek on the southeastern side of Cone-sus Lake (Livingston County). Baritized goniatites are com-mon in the Shurtleff Septarian Horizon (Bed 41a/5), whichcrops out on the northern bank just above creek level, 1.5 m(5 ft) below the Cashaqua/Rhinestreet contact, which is atapproximately 880 ft (268 m) elevation.

Locality 42*, Whetstone Brook.–A complete section ofCashaqua Shale is seen in Whetstone Brook above US Rte.20A, 2 mi (3.2 km) west of Honeoye (Ontario County).The section commences above the top of the Rock StreamSiltstone, which caps the first falls south (upstream) of thehighway. Marker black shale B is 3.7 (12 ft) above the con-tact and 6.7 m (22 ft) higher, baritized goniatites, includingManticoceras sinuosum clausium n. ssp. and Tornoceras unian-gulare obesum, are common in lenses within Bed 42/5, ahorizon of enormous concretions in the floor of the creek.Upstream, goniatites are less common in similarly large con-cretions of Beds 42/6-7, 0.46 m (1.5 ft) and 1.5 m (5 ft)above Bed 42/5. Bed 42/9 is a distinctive 25 mm (1 in)-thick brick-like limestone exposed just below the platformor floor of the stream at the base of the high falls. In the faceof the falls, a few goniatites were noted in the concretions ofBed 42/13, 8.2 m (27 ft) above Bed 42/9. Near the top ofthe falls, the Shurtleff Septarian Horizon (Bed 42/18) lies1.4 m (4.5 ft) below the base of the Rhinestreet Shale. Nogoniatites were recovered from the septarian nodules inplace, but a loose block with a rich fauna found near thebase of the Cashaqua section is believed to be from the hori-zon (locality 3004 of Kirchgasser, 1975).

Locality 43*, Briggs Gully.–The Cashaqua Shale is ex-posed in ravines above East Lake Road in the area of BriggsGully, on the southeastern side of Honeoye Lake (OntarioCounty). The lower Cashaqua is exposed in the third streamnorth of Briggs Gully and there a few goniatites were recov-ered from Bed 43/2, 3.0 m (10 ft) below marker black shaleB. In Briggs Gully, just above where outcrops begin, a rela-tively rich fauna of baritized goniatites was found in Bed43/3, a line of small concretions in the northern bank, 7.9m (26 ft) above the marker black shale and 2.4 m (8 ft)above a thin but prominent siltstone. Bed 43/3 is believedto be the type locality of Manticoceras sinuosum tardumClarke, 1898, which occurs with Probeloceras lutheri,Tornoceras uniangulare obesum, and Aulatornoceras. Near thetop of the section, fragments of baritized goniatites werenoted in the Shurtleff Septarian Horizon (Bed 43/13), 1.8m (6 ft) below the Cashaqua/Rhinestreet contact.


Locality 44*, Randall Gully.–Two major goniatite levelsare seen in the Cashaqua Shale of Randall Gully, 2.5 mi (4km) south of Bristol Center (Ontario County). The sectioncommences below Dugway Road at approximately 1,120 ft(341 m) elevation where a 0.1 m (4 in)-thick siltstone at thetop of a low falls marks the top of the Rock Stream Siltstone.A pair of thin siltstones mark the top of a 4 m (13 ft)-thickinterval of silty lower Cashaqua Shale and 2.4 m (8 ft) aboveis Bed 44/3, a 38 mm (1.5 in)-thick, discontinuous concre-tionary lens with clusters of baritized goniatites, mostlyManticoceras sinuosum clausium n. ssp. and Tornoceras unian-gulare obesum. Marker black shale B, with a rusty weatheringshale below, lies 3.2 m (10.5 ft) above Bed 44/3. Higher inthe section, a side stream enters from the north at approxi-mately 1,170 ft (356 m) elevation and 1.7 m (5.5 ft) aboveis a thin but prominent siltstone. The goniatitesM. sinuosumsinuosum, Probeloceras lutheri, and T. uniangulare obesum areparticularly common in Bed 44/4, a 50-100 mm (2-4 in)-thick argillaceous limestone lens that outcrops 2.3 m (7.5ft) above the siltstone. Above Bed 44/4, a few goniatites werenoted in the concretions of Beds 44/5 and 44/6 outcroppingjust downstream of the covered interval, which continues tothe base of the high falls, at the top of which is theCashaqua/Rhinestreet contact.

Locality 44a, Barnes Gully.–Two goniatite horizons werenoted in the discontinuous exposure of the Cashaqua Shalein Barnes Gully above Foster Point, Canandaigua Lake (On-tario County). In the southern branch of the gully, east ofNY Rte. 21, three small tributaries that join the creek fromthe southwest provide convenient geographic markers. Thelowest enters 1.1 m (3.5 ft) above the top of the waterfall atthe Rock Stream/Cashaqua contact. The second and largestside creek enters 2.9 m (9.5 ft) above the base of theCashaqua, and 0.76 m (2.5 ft) higher is a thin, dark gray,pyritic shale that is believed to be black shale B. The thirdside creek joins 0.9 m (3 ft) above this shale horizon, and inthe narrow gully of the tributary, 5.8 m (19 ft) above theconfluence, is a 6.4 cm (2.5 in) argillaceous limestone (Bed44a/2) with few but well-preserved goniatites, especiallyManticoceras sinuosum sinuosum and Probeloceras lutheri.Near the top of the section, at the point where exposuresbegin west of NY Rte. 21, goniatites (?Probeloceras lutheri)occur in the thin limestone beneath a bed of nodules (Bed44a/8), 4.3 m (14 ft) below the Cashaqua/Rhinestreet con-tact.

Locality 45, Hicks Point.–A complete sequence throughthe Cashaqua Shale is exposed in the ravine above HicksPoint, Canadaigua Lake, below NY Rte. 21, 2.3 mi (3.7 km)northeast of Bristol Springs (Ontario County). The sectioncommences at approximately 1,055 ft (322 m) elevationwhere a 15.2 cm (6 in)-thick siltstone at the top of a 4.6 m

(15 ft)-high waterfall marks the Rock Stream/Cashaqua con-tact. Goniatite fragments were noted in the nodular ParrishLimestone (Bed 45/2), 9.4 m (31 ft) above the contact and1.8 m (6 ft) above the point where the northern and south-ern branches join at approximately 1,080 ft (329 m) eleva-tion. Higher up, in the southern branch, 1.5 m (5 ft) abovethe Parrish Limestone, Manticoceras sinuosum, Probeloceraslutheri, and Tornoceras occur in the shales in the vinicity ofthe concretions of Bed 45/4 and in the shales of Bed 45/6,1.8 m (6 ft) above. A 0.9 m (3 ft)-thick interval of nodularshales begins 1.8 m (6 ft) above Bed 45/6, and 1.2 m (4 ft)and 2.4 m (8 ft) above the nodular interval are Beds 45/11and 45/13, respectively, thin limestones with goniatite frag-ments. In the upper part of the section, goniatites, includingProchorites alveolatus,M. sinuosum, P. lutheri, and Tornoceras,are common in the shales surrounding the nodules of Bed45/17 (locality 3014-17 of Kirchgasser, 1975), 1.5 m (5 ft)below the Shurtleff Septarian Horizon (Bed 45/18, withM.sinuosum sinuosum), which lies 2.3 m (7.5 ft) below theCashaqua/Rhinestreet contact; some of the septarian nod-ules at this locality contain abundant plant and fish frag-ments and, in places, baritized goniatites.

Locality 46, Cook Point.–A 20 m (65 ft)-thick section ofCashaqua Shale is found in the ravine above Cook Point,Canandaigua Lake, which crosses NY Rte. 21 at the edge ofBristol Springs (Ontario County). The succession com-mences above the high falls capped by a 0.13 m (5 in)-thicksiltstone that marks the Rock Stream/Cashaqua contact. Thefirst line of concretions above the contact is Bed 46/1 and afew goniatites are seen in the succeeding 1.5 m (5 ft) ofshale. They are more common, however, in the nodules ofBed 46/2, the Parrish Limestone, just below a covered inter-val 5.8 m (19 ft) above the base of the Cashaqua and in theshales surrounding a similar nodular horizon (Bed 46/3) 1.8m (6 ft) higher. A 0.15 m (6 in)-thick siltstone lies 1.2 m (4ft) above Bed 46/3 and a few goniatites were noted in theshales surrounding the concretions of Beds 46/7-8, 1.2 and2.0 m (4 and 6.5 ft) above the siltstone. TheCashaqua/Rhinestreet contact is at approximately 1,020 ft(311 m) elevation.

Locality 46a, Long Point.–Goniatites are common in the38 mm (1.5 in)-thick Parrish Limestone (Bed 46a/1) andcapping shales that crop out at approximately 980 ft (299m) elevation in the creek above Long Point, which crossesNY Rte. 21, 0.4 mi (0.6 km) southeast of Bristol Springs(Ontario County).

Locality 46b, Whiskey Point.–Molds of goniatites arecommon in the nodular shales of the 76 mm (3 in)-thickParrish Limestone (Bed 46b/1) in the ravine aboveWhiskeyPoint on the eastern side of Canandaigua Lake. The horizoncrops out at approximately 980 ft (299 m) elevation, 7.9 m


(26 ft) above the top of the Rock Stream Siltstone.Locality 46c, Snyder’s Gully.–At the southern end of

Canandaigua Lake, in Snyder’s or Snyder Gully, 0.4 mi(0.64 km) south of Woodville (Ontario County), a few go-niatites were recovered from the Parrish Limestone (Bed46c/1), a 50 mm (2 in)-thick nodular horizon exposed inthe creek bed above a covered interval 3 m (10 ft) above thetop of the high falls over the Rock Stream Siltstone. Thehorizon is approximately midway between NY Rte. 21 andGriesa Hill Road. This is locality 3015 of Kirchgasser(1975). The Middlesex Shale at this locality is the source ofNYSM 4077, the lectotype, selected herein, of Sandberge-roceras syngonum.

Locality 46d, Griesa Hill Road.–The Parrish Limestone(Bed 46d/1) crops out 3.4 m (11 ft) above the top of theRock Stream Siltstone in the third stream crossed by GriesaHill Road north of the intersection with NY Rte. 21, 3 mi(4.8 km) north of Naples (Ontario County). This is locality3009 of Kirchgasser (1975).Manticoeras sinuosum sinuosumand Probeloceras lutheri are relatively common in the ParrishLimestone here and at most localities in the area nearNaples.

Locality 46e, Bristol Springs Road.–The Parrish Lime-stone (Bed 46e/1) forms a ledge in a tributary to NaplesCreek that crosses NY Rte. 21, 0.9 mi (1.4 km) northeast ofthe intersection of the highway and the Bristol SpringsRoad, Naples (Ontario County).

Locality 46f, Bristol Springs Road Gully.–In the lowerpart of the narrow gully above the intersection of NY Rte.21 and the Bristol Springs Road, Naples (Ontario County),the Parrish Limestone (Bed 46f/1) is 76 mm (3 in) thickand lies 2.7 m (9 ft) above the top of the Rock Stream Silt-stone.

Locality 47*, Conklin Gully.–One of the most completeand accessible sections of Cashaqua Shale in the Naples areais the 15 m (50 ft)-thick sequence above the first falls inConklin Gully (formerly Parrish Gully), approximately 400yd (0.4 km) southeast of the intersection of Rumpas Hill(Parish) Road and NY Rte. 245, northeast of Naples (On-tario County). This is the type locality of the Parrish Lime-stone (Bed 47/2), the “Goniatite Concretionary Layer” ofClarke (1885), which is the 10.2 cm (4 in)-thick nodularlimestone outcropping along both banks of the creek, 2.1m (7 ft) above the Rock Stream/Cashaqua contact (Kirch-gasser, 1965, et seq.).Manticoceras sinuosum sinuosum is com-mon and Probeloceras lutheri and Tornoceras occur. A similarbut unfossiliferous horizon lies 2.1 m (7 ft) above Bed 47/2and a few goniatites were noted in the shales beneath thesmall concretions of Bed 47/4, 0.9 m (3 ft) higher. In thedark shales at the top of the section, baritized goniatite frag-ments were noted in Bed 47/12, the easternmost locality of

the Shurtleff Septarian Horizon, 0.9 m (3 ft) below the baseof the Rhinestreet Shale.

Locality 47a, Rumpus Hill.–Goniatites are also commonin the Parrish Limestone (Bed 47a/1) in the tributary to theWest River on Rumpas Hill, which crosses NY Rte. 245 ap-proximately 5 mi (8 km) northeast of Naples (OntarioCounty). The gully is 0.1 mi (0.16 km) northeast of theturnoff leading to Clark Gully, which is on the opposite sideof the valley. The horizon consists of 12.7 cm (5 in) of nodu-lar shale and lies 0.9 m (3 ft) above the top of the RockStream Siltstone.

Locality 47b, Clark Gully.–On the eastern bank of ClarkGully, on the southern side of South Hill, 5.3 mi (8.5 km)northeast of Naples (Ontario County), goniatites are com-mon in the 0.1 m (4 in)-thick Parrish Limestone (Bed47b/1) that crops out 1.8 m (6 ft) above the top of the RockStream Siltstone.

Locality 47c, Lee Cemetery.–A few goniatites (Mantico-ceras sinuosum and Probeloceras lutheri) were recovered fromthe 0.13 m (5 in)-thick Parrish Limestone (Bed 47c/1) thatcrops out 1.2 m (4 ft) above the Rock Stream Siltstone in thegully crossing NY Rte. 245, 0.1 mi (0.16 km) northeast ofLee Cemetery and 5.7 mi (9.2 km) northeast of Naples (On-tario County).

Locality 47d, School No. 7, Middlesex.–Manticocerassinuosum sinuosum is particularly common in the ParrishLimestone (Bed 47c/1), cropping out in a stratigraphic set-ting similar to Loc. 47b, in the large ravine 0.1 mi (0.16km) northeast of the site of old School #7 onWest Avenue,1.7 mi (2.7 km) southwest of Middlesex (Yates County).

Locality 47e, Lower East Hill Road.–In the gully be-tween NY Rte. 245 and Lower East Hill Road, 0.9 mi (1.4km) south of Middlesex (Yates County), the nodular ParrishLimestone (Bed 47e/1) is poorly exposed but has yielded afew goniatites.

Locality 47f, School No. 2, Italy Valley.–In the lowerpart of the creek that crosses the Italy Valley Road at the siteof old School #2, 3.2 mi (5.1 km) north of Italy (YatesCounty), the Parrish Limestone (Bed 47f/1) is a 0.1 m (4in)-thick nodular shale withManticoceras sinuosum, cappedby shales with abundant auloporid corals, which crops outopposite the school site and lies 0.9 m (3 ft) above the RockStream Siltstone.

Locality 48*, Wagener Glen.–A condensed, 4.3 m (14ft)-thick section of Cashaqua Shale is exposed in WagenerGlen (West Branch of Keuka Lake) above the first falls westof the roadbridge, 0.5 mi (0.8 km) north of Pulteney(Steuben County). There the 0.28 m (11 in)-thick ParrishLimestone (Bed 48/1) is only 76 mm (3 in) above the 0.3 m(1 ft) ledge of siltstone that marks the top of the RockStream Siltstone. No goniatites have been collected from the


Parrish Limestone, but Probeloceras lutheri is common in abioturbated shale (Bed 48/2) beneath a 76 mm (3 in)-thickblack shale 0.3m (1 ft) above. A sequence of siltstones, siltygray shale, and black shale continues upstream to theCashaqua/Rhinestreet contact.

Locality 49, Bobell Hill.–At Bobell Hill Quarry, 4 mi(6.4 km) southwest of Coventry (Chenango County), J.Harrington collected a specimen ofManticoceras that is theeasternmost known occurrence of the genus in New YorkState. The locality is just south of Schaffer Road, at an alti-tude of approximately 1,540 ft (470 m), west of Bobell Roadand Bobell Hill. The stratigraphical equivalence of the levelwith horizons to the west is uncertain. The facies in thequarry is that of Cypricardella and the possible equivalent isthe Enfield. The level would seem to fall within the SonyeaGroup.

WEST FALLS GROUP

This group comprises the Rhinestreet and Angola Shale For-mations of New York State west of Varysburg (WyomingCounty), and their more clastic easterly equivalents. Raisedfrom formation status (Rickard, 1964; Oliver et al., 1969)to group status by Rickard (1975), the division is defined asthose strata between the basal black shales of the RhinestreetShale and the base of the black shales of the Pipe Creek ShaleFormation (Text-fig. 2).

Rhinestreet ShaleThis is the thickest of the Upper Devonian black shale in-tercalations in western New York. It was recognized by Hall(1843) as the “upper black shale.” The unit was defined byClarke (1903: 23) and by Luther (1903) under the nameRhinestreet Shale or “black shale” and later “bituminousshale” (Clarke, 1904: 213) or “second black band” (Clarke,1898: 106). The division was named for exposures nearRhinestreet Road, Naples, a sequence that represents onlythe early part of the unit farther west.

In the detailed study of the West Falls Group by Pepperet al. (1956), the Rhinestreet Shale was measured as 59.4 m(195 ft) thick at Eighteenmile Creek, thinning eastwardlyto 20.7 m (68 ft) thick at the Mount Morris Dam outcropon the Genesee River, and to 4.9 m (16 ft) thick in UrbanaGlen in Steuben County, but it was recognized that this re-duction resulted from facies changes. The top of theRhinestreet Shale was taken at the bed named the “ScraggyBed” by Luther (1903: 1022-1023), a distinctive bed ofpyritic pustulate concretions. Pepper et al. (1956) were un-able to trace this bed east of Cazenovia Creek, but we wereable to trace the bed of black shale that overlies it muchmore extensively (Text-figs 13-14) and doubtless additionalwork will carry it yet farther, because the black shale forms

a better unit for definition of the top of the RhinestreetShale.

The beds of the lower two thirds of the Rhinestreet Shalein the western half of Erie County are “composed largely ofbrownish-black and petroliferous shale in which an upwardgradation from black into dark brown or very dark gray atintervals of 2 inches (5.1 cm) to medium dark gray tomedium-gray shale in beds from an inch (2.54 cm) to 5 ft(1.52 m) thick” (Pepper et al., 1956).

It is in the upper part of the Rhinestreet Shale in thesame area that the microcycles so characteristic of the over-lying Angola Shale are progressively developed. The cyclesare initiated with black or brown shale, the upper part ofwhich shows evidence of burrows as it passes rather suddenlyinto medium-gray shale, the upper part of which shows thedevelopment of calcareous, and often septarian, concretions.It is the concretionary levels that frequently produce well-preserved goniatite faunas, but mostly only in the upper partof the formation (Text-fig. 13). The most productive hori-zon is the Relyea Creek Horizon at Relyea Creek (Loc. 60/3)in Wyoming County, and correlative or nearly correlativehorizons at Cazenovia Creek (Loc. 54/1), Varysburg (Loc.57/1, 2), Johnson Creek (58/1), Stony Creek (61/1), andKennedy Gulf (63/3), which have yielded Sphaeromantico-ceras oxy (Clarke, 1897), S. aff. oxy, S. rhynchostomum(Clarke, 1898), Manticoceras lamed (G. & F. Sandberger,1850), Carinoceras vagans (Clarke, 1898), Playfordites cf. tri-partitus (G. & F. Sandberger, 1850) (Loc. 58/1 only), Lin-guatornoceras aff. linguum (G. & F. Sandberger, 1851), andTornoceras cf. typum (G. & F. Sandberger, 1851) (Text-figs13, 61).

No substantial effort has been made in this work to es-tablish a detailed correlation of the lower Rhinestreet Shale(Text-fig. 11). This part of the sequence is not rich in goni-atites, and they are known from only a few localities.

At Buck Run Creek (Loc. 65/1), for example (Text-fig.11), a fauna with Prochorites aff. alveolatus occurs associatedwith a fossil-log concretionary bed 12.7 m (41.7 ft) abovethe base of the Rhinestreet Shale. The Sturgeon Point (Loc.50) Lake Erie area is recorded as having yielded manticocer-atids (Clarke, 1899c), but the exact source has not been lo-cated. Large concretions to 2.4 m (8 ft) across withmanticoceratids occur in the vinicity of the northernpromontory of Grandview Bay, Lake Erie, at a level thatmust be approximately 10.7 m (35 ft) below the ScraggyBed, that is, quite high in the sequence. The sections illus-trated for the lower Rhinestreet Shale and the sedimentarysetting show that detailed lithostratigraphic work would en-able correlation of the thin leaves of black shale that pro-gressively break away from the basal black shales as one goeseastward.


The sedimentary microcycles of the upper RhinestreetShale are traceable from near Lake Erie eastward to KennedyGulf (Loc. 63) in Wyoming County (Text-fig. 13). As Pep-per et al. (1956) noted, the Scraggy Bed is soon lost, but theoverlying black shale is a reliable marker for the top of theRhinestreet Shale.

Luther (1903: 1023) noted the Scraggy Bed at lake level

at Fox’s Point on the southern side of Dibble Bay, Lake Erie,but it is now best exposed in Big Sister Creek (Loc. 73, Text-fig. 15) and Cazenovia Creek (Loc. 54, Text-fig. 13). Cor-relation of this level eastward shows that when plotted onthe chart of Pepper et al. (1956), it lies 67 m (220 ft) abovethe base of their Kennedy Gulf section (no. 8) at the level ofthe uppermost black shale that they indicate. Thus, the cor-

Text-fig. 11. Sections of the lower Rhinestreet Shale and its equivalents at Tannery Brook, Attica (Wyoming County), Buck Run Creek,Mount Morris (Livingstone County), and Parrish or Conklin Gully (Yates County, near Naples, Ontario County). Note: for Loc. 69, readLoc. 65a.


relation here established shows that most of the GardeauShale and the Grimes Siltstone and Hatch Shale of the re-gion east of Varysburg is the time equivalent of the Lake ErieRhinestreet Shale at Lake Erie (Text-fig. 3).

Much effort was expended in trying to extend this cor-relation into the Genesee Gorge (Loc. 86) section nearMount Morris (Text-fig. 3). The level seems to be well abovethe Table Rock Sandstone of Hall (1843: 391), Luther(1903: 1006), and Chadwick (1933: 96), that is the bed thatforms the top of the Lower Falls (or Lower Portage of theGenesee River (Text-fig. 12), but the exact correlation hasnot been established. Pepper et al. (1956) did not mark theTable Rock Sandstone on their stratigraphical column (no.12) but they described it and it is the unit that they showed51 ft (15.5 m) above the base on their column no. 12 be-cause they indicated that this level is 395 ft (120 m) abovethe top of the Rhinestreet Shale in the Genesee River. Thetop of the Middle Falls lies 190 ft (58 m) higher and the topof the Upper Falls seems to lie immediately underneath theNunda Sandstone on their chart.

There are a few type specimens of goniatites from thetime equivalents of the Lake Erie Rhinestreet Shale in theGenesee Valley region. The source of the syntypic specimenof Sphaeromanticoceras oxy (see Clarke 1898, 1899c, 1899a,1899b: 78, pl. 2, figs 5-6; NYSM 3746) was given as “fromthe soft shales, 150 ft below the Portage sandstone, at theLower Portage Falls on the Genesee river.” Accepting the lo-cation rather than the elevation as being the more reliable,this level would be equivalent to the upper part of theRhinestreet Shale of Lake Erie and not the Angola Shale(Miller, 1938: 95). The other specimen originally figured byClarke (1898, 1899a, 1899b: text-fig. 61; NYSM 12305/4)was said to be from Stony Brook Glen (Steuben County;Loc. 87), just south of Dansville (Livingston County). Thesection there was illustrated by Pepper et al. (1956, sectionno. 18) and, accepting their correlation of the sequence fromthe uppermost Gardeau Shale to upper Nunda Sandstone,the specimen is most likely to have come from equivalentsof the Angola Shale of Lake Erie.We remeasured this sectionto establish that the base and top of the main falls lie at 61m (200 ft) and 74.7 m (245 ft), respectively, above the baseof their section 18 and wholly within their West Hill Mem-ber.

Localities.–Locality numbers with asterisks have sections il-lustrated in Text-figs 11-13. The localities given here reallycomprise two groups. Localities 50-55 and 65 comprise lev-els generally referred to the Rhinestreet Shale. But most ofthe other localities represent horizons that are correlated herewith the Rhinestreet Shale near Lake Erie on the basis ofbed-by-bed correlations (Text-fig. 12), but which, litholog-

Text-fig. 12. Section at the Lower Portage Falls on the GeneseeRiver, Livingston County.


Text-fig. 13. Sections of the upper Rhinestreet Shale equivalents between Cazenovia Creek, Erie County, and Kennedy Gulf north ofWarsaw, Wyoming County, with the locality numbers used in the text.



ically, are referred to the Gardeau Shale farther east.Locality 50, Sturgeon Point.–An old locality from which

manticoceratids have been reported (Clarke, 1899c) is Stur-geon Point, on Lake Erie Shore (Erie County) at the north-ern end of Grandview Bay. Giant septarian nodules to 2.4 m(8 ft) across, exposed near Sturgeon Point, containMantic-oceras sp. This locality might lie approximately 10.7 m (35ft) below the black marker band above the Scraggy Bed atPoint Breeze to the south (see Addendum).

Locality 51, Big Sister Creek.–Along Big Sister Creek,near Angola (Erie County), the upper Rhinestreet sectioncontinues downward from the Angola and the relevant hori-zons are illustrated there (Loc. 73, Text-fig. 15). There aretwo levels of concretions below the Scraggy Bed that haveyielded goniatites: Bed 51/b at 0.46 m (1.5 ft) below, andBed 51/a at 1.37 m (4.5 ft) below.

Locality 52, Eighteenmile Creek.–The good sectionalong Eighteenmile Creek (Erie, Chautauqua, and Catta-raugus counties; Grabau 1898-1899; Pepper et al., 1956:section 1) has not been systematically searched for goni-atites.

Locality 53*, Hampton Brook.–The Hampton Brooksection south of Hamburg (Erie County; Loc. 74, Text-fig.15), illustrated for the Angola Shale equivalents, continuesdown into the upper Rhinestreet, but faunas have not beencollected.

Locality 54*, Cazenovia Creek.–The upper part of theRhinestreet Shale in Cazenovia Creek (Erie County) con-tains concretions similar to those in the overlying AngolaShale (Loc. 75, Text-fig. 15). Manticoceras lamed, Sphaero-manticoceras rhynchostomum, and Tornoceras cf. typum werecollected from the concretions of Bed 54/1, 2.4 m (8 ft)below the basal black shale marker of the Angola Shale, and3.5 m (11.5 ft) above a prominent bed of septarian concre-tions (Text-fig. 13).

Locality 55*, Hunter Creek.–The top of the Rhinestreetis exposed 200 yd (183 m) east-northeast of a bridge east ofColegrove at approximately 1,033 ft (315 m) altitude inHunter Creek (Erie County), and a long section below is il-lustrated here (Text-fig. 13). The main collection here (Loc.55) came from a loose block collected below the ScraggyBed. This was probably derived from the Angola Shale. Oth-erwise no goniatites were collected from the section.

Locality 56, Sheldon Creek.–North of Strykersville, inSheldon Creek (Wyoming County), at the very base of thesection exposed above glacial debris, a level of concretions(Bed 56/a) 3 ft (0.9 m) below the marker black shale of theAngola, at approximately the level of the Scraggy Bed,yielded Tornoceras sp. (Loc. 78, Text-fig. 15).

Locality 57*, Varysburg.–The section along TonawandaCreek at Varysburg (Wyoming County) commences approx-

imately 120 yd (109 m) upstream of the US Rte. 20A bridgewest of the town (Loc. 57, Text-fig. 13). A continuation up-ward of the section is illustrated inText-fig. 15 (Loc. 79/80).At the lower locality Bed 57/1 is a horizon of small concre-tions below a 12.7 cm (5 in)-thick siltstone. Bed 57/2, justover 2.1 m (7 ft) higher, is a fossil-log concretion horizonwith a log of fossil wood, 0.66 m (2.2 ft) below a 0.13 m (5in) black shale. The goniatite fauna from this level is excep-tionally rich and includes Carinoceras vagans, Manticoceraslamed, Sphaeromanticoceras rhynchostomum, and Lingua-tornoceras aff. linguum. Localities hereabouts were knownto Williams (1887) and formed his locality 472.

Locality 58*, Johnson Creek.–The upper part of theRhinestreet Shale is exposed above a cabin, 0.5 mi (0.8 km)west-northwest of Earls, in Johnson Creek (WyomingCounty), 200 yd (180 m) west of a disused railroad. Bed58/1 (the Relyea Creek Horizon) lies 0.66 m (2.2 ft) belowthe base of a 0.18 m (7 in) black shale above a thin siltstonethat forms the foot of the main falls. The rich fauna of Bed58/1 includes Carinoceras vagans, Manticoceras lamed,Sphaeromanticoceras rhynchostomum, Playfordites cf. triparti-tus, and Linguatornoceras aff. linguum. The upward exten-sion of this section (Loc. 81) is given in Text-fig. 15.

Locality 59a*, Tannery Brook.–A section in the lowerRhinestreet Shale continues upward from the UpperCashaqua Shale (Kirchgasser, 1967: 138, loc. 7), 1.2 mi (2.3km) west of Attica (Wyoming County; Text-fig. 11). Thebase of the Rhinestreet Shale is at approximately 1,100 ft(335 m) elevation. No goniatites were found in theRhinestreet Shale at this locality.

Locality 60*, Relyea Creek.–Near South Warsaw in Re-lyea Creek (Wyoming County) is a locality known to Clarke(1899c) and mentioned by him in his Guide and possiblyLuther (1903: 1012) under the name of Gibson’s Glen. Thesection given here (Text-fig. 13) commences 500 yd (460m) west of NY Rte. 19 at South Warsaw at approximately1,100 ft (335 m) altitude, above which a farm track crossesthe creek at a falls over a thick black shale. Upstream a rockpool forms the most convenient marker for the goniatite-rich levels. Bed 60/2 is approximately 0.9 m (3 ft) above thepool surface, and 0.46 m (1.5 ft) above is a 12.7 cm (5 in)-thick siltstone that marks the top of a small falls. Bed 60/3,the Relyea Creek Horizon, is above a 0.23 m (9 in)-thickblack shale, 0.84 m (2.75 ft) higher and 5.6 m (18.4 ft)below the Rhinestreet/Angola Shale contact. The rich faunaof Bed 60/3 includes Sphaeromanticoceras oxy, S. aff. oxy,Carinoceras vagans,Manticoceras sp., S. rhynchostomum, andLinguatornoceras aff. linguum. Bed 60/4 contains Mantico-ceras lamed.

Locality 61*, Stony Creek.–At Stony Creek, Warsaw(Wyoming County), a good and complete section of the


equivalents of the upper Rhinestreet is exposed (Text-fig.13). The locality was known to Luther (1903: 1012) as FallBrook. The section commences where exposures begin,1,400 yd (1.3 km) southwest of the town center. The onlygoniatites located (Sphaeromanticoceras rhynchostomum andTornoceras cf. typum) are from Bed 61/1, a level apparentlyequivalent to the productive interval at Relyea Creek. Bed61/1 is best approached from the railroad crossing, 2,200yd (2 km) southwest of Warsaw at which the upperRhinestreet Shale and lower Angola Shale equivalents are ac-cessable high in the face of the spectacular waterfall (Text-fig. 13, Loc. 61; Text-fig. 14, Loc. 83). The marker blackshale at the base of the Angola Shale equivalents is 10.9 m(35.7 ft) below the base of the 15 cm (6 in)-thick siltstoneon which the railroad bridge rests, and 7.3 m (24 ft) belowthe base of a 0.3 m (1 ft)-thick siltstone that caps the mainfalls. There is a 12.7 cm (5 in)-thick siltstone 6.8 m (22.4 ft)lower, and the concretions of Bed 61/1 are within a 0.81 m(2.7 ft) interval below a 15.2 cm (6 in) black shale.

Locality 62, Warsaw.–A roadcut along US Rte. 20A,west of Warsaw (Wyoming County) yielded goniatites 107yd (98 m) downhill from the intersection with Buffalo Av-enue (the old US Rte. 20A) at an altitude of approximately1,230 ft (375 m).

Locality 63*, Kennedy Gulf.–There is a fine section inKennedy Gulf, a tributary of Little Tonawanda Creek, 0.75mi (1.2 km) south of Dale (Wyoming County). The section(Text-fig. 13) commences 14 m (6 ft) above where a sidecreek enters at approximately 1,273 ft (388 m) altitude. Theupper section can be reached from a north-south road lead-ing to Vernal. Convenient markers in the section are few,and none are marked on the section of Pepper et al. (1956:section 8). The top of the section given here is the crest ofthe main waterfall in the section most easily reached fromthe Vernal road. This appears to be a thin sandstone unitthat Pepper et al. (1956) marked at 267 ft (81 m) below theblack Pipe Creek Shale at the top of their section. Themarker black shale taken as the base of the Angola Shaleequivalent lies 5.8 m (19 ft) below and appears to be thehighest black shale they showed. At 34 m (37 ft) below thecrest of the main waterfall is a small falls over a 0.18 m (7in)-thick siltstone, and the best fauna collected (withMan-ticoceras lamed and Sphaeromanticoceras rhynchostomum) isfrom the concretions of Bed 63/3, the probable RelyeaCreek Horizon, which outcrops 2.6 m (8.5 ft) lower, imme-diately above the top of the first falls encountered workingup the section. The most distinctive band below, just abovethe base of the section given here, is a 12.7 cm (5 in)-thickblack shale with a 50 mm (2 in)-thick siltstone in the mid-dle, a horizon seen also in Stony Creek. Bed 63/1, withAulatornoceras aff. eifliense, is the concretionary level in the

microcyclothem above this, and Bed 63/2 is at the top ofthe next cycle above which the creek floor forms a long gully.

Locality 65*, Buck Run Creek.–Ten miles (16 km)southeast of Mount Morris (Livingston County), Buck RunCreek exposes the Rhinestreet and Cashaqua Shale andlower units. The section given here (Text-fig. 11) is in theupper part of the section just downstream from where theSwanson Road Bridge crosses the creek. A 40 cm (1.3 ft)-thick black shale is the top bed marked on the section givenhere (Text-fig. 11, Loc. 65). Bed 65/1, a fossil-log locality,has the goniatite Prochorites aff. alveolatus. The base of theRhinestreet lies 12.8 m (42 ft) below Bed 65/1 at an altitudeof approximately 755 ft (230 m). The higher horizonsmarked (Beds 65/2 and 65/3) were collected for conodontsby J. W. Huddle, but goniatites have not been found inthem.

Locality 65a*, Parrish Gully.–At the Canandaigua Lakemeridian (Text-fig. 3), a section in the lower RhinestreetShale equivalents continues upward from the CashaguaShale in Parrish or Conklin Gully (Loc. 47) in Yates County,northeast of Naples (Ontario County; misnumbered 69 inText-fig. 11). No goniatites were found in the RhinestreetShale equivalents at this locality.

Equivalents of the Sonyea andWest Falls Groups Southand Southeast of Ithaca

There is a remarkably confined area in which the generaSandbergeroceras, Wellsites, and Schindewolfoceras occur inthe area south and southeast of Ithaca (Tompkins County;Text-fig. 14). The precise horizons represented by the vari-ous localities is a matter for individual discussion. Manyseem relatable to the tongues of black shale that Sutton(1963) and Sutton et al. (1962) traced in the area (Rickard,1964, 1975), that is, for Middlesex Shale equivalents theMontour and Sawmill Creek Shales, and for the Rhinestreetequivalents the successive sequence of the Moreland, DunnHill, Roricks Glen, and Corning Shales. These tongues haveyet to be traced westward with precision into the standardsequence.When this is done, it could be that these goniatiteswill be found to be more widely distributed. At present, itis only the Sandbergeroceras faunas that are known fartherwest, as remarked earlier, in the Naples and Honeoye areaswithin the Middlesex and Cashaqua Shales. There is also apossible record in the Tygart Valley (Randolph County),Virginia (Tilton, 1929; Miller, 1938: 180).

The records seem to comprise a sequence in which (1)Sandbergeroceras syngonum occurs in equivalents of the Mid-dlesex Shale, (2) Sandbergeroceras? enfieldense n. sp. occursin equivalents of the upper Cashaqua Shale and lowerRhinestreet Shale, (3) Wellsites williamsi (Wells, 1956) andW. tynani House & Kirchgasser, 1993, occur in lower


Rhinestreet equivalents, and (4) Schindewolfoceras chemu-ngensis Vanuxem, 1842, S.? equicostatum (Hall, 1874), andS.? aff. equicostatum occur in the equivalents of the middleor upper Rhinestreet Shale.

Localities.–It should be noted that although these localitiesare all included in the West Fall Group section, some lie inequivalents of the Sonyea Group of the western part of thestate.

Locality 64, Harford Mills.–Harford Mills (CortlandCounty) lies 15 mi (24 km) east of Ithaca (TompkinsCounty). One of H. S. Williams’s (1913) recurrent Tropi-doleptus Zone localities (Hd 12G/1b) provided Sandbergero-ceras syngonum said to be from the Enfield Shale. The localityis southwest of Harford Mills (Tioga County) at approxi-mately 1,270 ft (387 m) altitude, along the stream that flowseastward fromWest Hill and crosses NY Rte. 38, 1.2 mi (2km) south of the county line. We also found specimens ofSandbergeroceras (NYSM 16577 and 16578) in this gully ata similar level. L. V. Rickard suggested (pers. comm., ca.1985) that the level is below that of the Moreland Shale andprobably just above the Sawmill Creek Shale, that is, anupper Middlesex Shale equivalent.

Locality 66, Hungerford Quarry.–The University quar-ries lie approximately 2.9 mi (4.6 km) east-southeast of theCornell University campus, in Ithaca (Tompkins County),and south of Ellis Hollow Road. There are several quarriesapproximately along the 1,200 ft (366 m) contour. The olduniversity quarry, or Hungerford Quarry, is the more west-erly, and best approached from Ellis Hollow Road; thenewer quarries are on either side of the road over the hill, 1.5mi (2 km) south of Turkey Hill. One large specimen ofSandbergeroceras? enfieldense n. sp., collected by K. Caster in

1934, is the holotype (NYSM 12028). The specimen is re-ported as having come from a layer containing the spongesIthacadictya and Protoarmstrongia. The “Endfield” here isprobably the equivalent of the Cashaqua Shale of the west(Rickard, 1964) and the black shale at the top of the quarriescould be the equivalent of the Rhinestreet Shale.

Locality 67, Fairfield Forest.–From a locality 1.8 mi (2.9km) southwest of Speedsville (Tioga County), 0.7 mi (1.1km) east of Fleet Road on States Lands Road at an elevationof approximately 1,640-1,660 ft (500-506 m), a specimenof Schindewolfoceras? aff. equicostatum was collected by J. W.Wells in 1963 (his locality 389) from a small quarry near ayouth camp in Fairfield State Forest. This appears to be thesmall quarry marked on the topographic map. We locatedno goniatites there. The horizon was recorded as being fromthe Upper Cayuta Shale (Chesney Formation) approxi-mately 400 ft (122 m) above a Tropidoleptus level. L. V.Rickard suggested (pers. comm., ca. 1985) that the levelcould be between the Moreland Shale and the Roricks GlenShale, that is, a mid-Rhinestreet Shale equivalent.

Locality 68, Bald Mountain.–The source of the holotypeof Wellsites williamsi is from beside a track 2 mi (3.2 km)southeast of Brooktondale, 7 mi (11.3 km) southeast ofIthaca, and 1.7 mi (2.7 km) northeast of White Church(Tioga County), and 1.2 mi (1.9 km) northwest of BaldMountain at an elevation of approximately 1,770 ft (539m). This is the specimen referred by Wells (1956b) to Belo-ceras. It was recorded as coming from the “lower part of theCayuta Shale (Chemungian Stage, upper Senecan Series),approximately 150 ft (46 m) above the Thiemella (?) danbyizone (= base of Cayuta)” (Wells, 1956b: 751). The levelwould be above the Moreland Shale, but not far above. Thislocality seems to be from along Bald Hill School Road–Leonard Road Junction (Loc. 68). Another locality, southof the junction, is Caroline (Loc. 94), Bald Hill, town ofCaroline, which is the source of Sandbergeroceras? enfieldensen. sp. (NYSM 12027).

Locality 69, Owego.–From the general area of Owego(Tioga County), there are some old records of goniatites.The locality of the holotype of Schindewolfoceras chemu-ngense (Vanuxem, 1842) (NYSM 4073) was given asOwego, and a subsequent label added “Cayuta Shale”; anold label gave the number 2 (?8) and “-hem Quarry, nearOwego.” The identification of this quarry has not been de-termined but there have long been quarries on the hillsidejust north of Owego. L. V. Rickard suggested (pers. comm.,ca. 1985) that they must be at levels equivalent to theRhinestreet Shale of the west; the contact between the Ror-icks Glen, Beers Hill, and Moreland Members is below thetop of East Beecher Hill here. A Manticoceras sp. specimen(CU G54) in the CU collections is listed as from “locality

Text-fig. 14. Map of localities withTriainoceratidae in rocks equiv-alent to the Sonyea and West Falls groups in central New YorkState and the locality numbers used in the text. Geology fromRickard & Fisher (1970).



29” on the southeastern side of Beecher Hill.We found horizons with goniatites on Mountain Road

or East Beecher Hill Road, which leaves Owego just northof the railroad crossing of NY Rte. 96. The entrance of Ever-green Cemetery is along the road and at 0.33 mi (0.5 km)farther up the hill, a 2 m (6 ft) bed of siltstone has largeslump structures; higher up, a fallen block yielded Mantic-oceras (NYSM 16579). The sandstones and siltstones thereare commonly full of cyrtospiriferids, productids, and bi-valves.

Locality 70, Elmira.–The holotype of Wellsites tynani(SUI 42318) was collected by Mr. M. C. Tynan of the Uni-versity of Iowa in 1978 frommaterial derived from a roadcutfor US Rte. 17, which cuts through the Moreland Shale, 0.1mi (0.16 km) east of the crossing of NY Rte. 116 and Wa-tercure Road, Elmira (Chemung County), and just belowthe East Church Street Quarries (which formed locality 7of Woodrow and Nugent’s itinerary for the thirty-fifth An-nual Meeting of the New York Geological Association;Coates, 1963) at which the junction between the Millportand Dunn Hill units was identified. The horizon of the lo-cality would seem to be below 910 ft (277 m) in altitude, thefigure quoted byWoodrow and Nugent. The completion ofthe US Rte. 17 bypass has led to the sections there beingovergrown or obscured, but there are fine sections a little tothe south at an easterly exit to Church Street, Elmira, wherepoorly preserved goniatite remains were noted. There is aproblem as to the horizon because Sutton et al. (1962: 392)stated that the type section of the Roricks Glen is at a scoutcamp at the southern end of Elmira, and that is an upperRhinestreet unit. It remains to be investigated whether thestratal dip would carry the supposed Moreland Shale belowthe type section of the Roricks Glen.

Locality 94, Bald Hill, Caroline.–A specimen of Sand-bergeroceras? enfieldense n. sp. collected by S. C. Hollister isin the CU collections (1071K; plastotype NYSM 12027). Itcame from Bald Hill, town of Caroline (Tompkins County),approximately 0.5 mi (0.8 km) south of Loc. 68 at an alti-tude of between 1,700 and 1,800 ft (518 and 549 m). Thislevel is not far above the Moreland Shale.

Locality 95, North Chemung.–A single poor specimenof Schindewolfoceras? sp. with Tropidoleptus in the CU collec-tions (Bartwick collection, Locality 48 or 29?) is recordedfrom the Chemung Group of authors between Chemungand North Chemung (Chemung County). No further infor-mation is available on this locality.

Locality 96, Van Etten.–A single triainoceratid collectedin 1882 by H. S. Williams (his locality LXII), CU 3308, islabeled as from Van Etten (Chemung County; WaverlyQuadrangle). No further information is available on this lo-cality.

Angola ShaleThe Angola Shale was originally named for the sequence ofgrey shales that overlies the Rhinestreet at approximately thevillage of Angola (Erie County; Clarke, 1903: 24; Luther,1903: 1019). The thickness assigned to the unit by Pepperet al. (1956) was 68.6 m (225 ft) in Hampton Brook (Loc.74) and 71.6 m (235 ft) in Cazenovia Creek (Loc. 75) (Text-fig. 15).

Pepper et al. (1956) demonstrated that the Nunda Sand-stone and West Hill Flags intercalate from the east in theupper parts of the equivalents of the Angola Shale (see alsoBaird & Jacobi, 1999). They took the 0.6 m (2 ft)-thick silt-stone bed at West Falls on Cazenovia Creek, approximately20.1 m (66 ft) below the top of the Angola Shale there, asthe westernmost edge of the Nunda Sandstone (this unitcrops out just downstream from the bridge at West Falls atan altitude of 920 ft or 280.4 m). In this part of the section,we found an underlying group of three black shales (infor-mally herein called “The Trinity”) that enabled correlationeastward to Relyea Creek (Loc. 82). The correlation of thelowest black shale of “The Trinity” is shown in Text-fig. 15.Associated with these black shales and siltstones are a num-ber of goniatite horizons both at the localities shown (Text-fig. 15) and others.

The sedimentary microcycles of the Angola Shale con-tinue those of the Rhinestreet, and in the lower part theyare developed particularly well as a group of units that canreadily be traced well to the east (Text-fig. 15). Thin silt-stones progressively enter eastward and up the succession.One such thin turbidite crops out at the mouth of FarnhamCreek at Point Breeze (Loc. 72) Lake Erie Shore (above Bed72/6, Text-fig. 15) and can be traced for approximately 45mi (72 km) to Johnson Creek (Loc. 81). The concretionaryunits in the lower part of the Angola Shale are particularlyrich in well-preserved goniatites.

The Angola Shale is the source of some of the primarycollections. The type material of Sphaeromanticoceras rhyn-chostomum (Clarke, 1898: 65, pl. 4, figs. 6-13, pl. 5) in-cludes material from Big Sister Creek (Loc. 73) presumablyfrom the localities later indicated by Clarke (1899c: 107) asnear the crossing of the Lake Shore railroad and the creek;other material from Clarke (NYSM 3794-6) is labeled onlyas Angola. The type material of Carinoceras sororium(Clarke, 1898: 75, pl. 4, figs. 1-5, text-fig. 55) was said tocome from “the vicinity of Angola, on the Lake Erie shoreand along Big Sister and Farnham creeks, Erie County.” Thisagain would indicate the concretionary levels low in the An-gola Shale but, as will be seen (Text-fig. 15), almost all theconcretionary levels of the microcyclothemic units here yieldgoniatites so that the exact level is likely to remain uncertain.The most productive horizon is the Point Breeze Goniatite

Text-fig. 15. Sections of the lower Angola Shale equivalents between Lake Erie and Warsaw, Wyoming County, with the locality numbersused in the text. Note: For Loc. 75, read 1 mi NE of Griffins Mills; for Loc. 80, read Loc. 79.



Bed, which has yielded Sphaeromanticoceras oxy, S. rhyncho-stomum, Carinoceras vagans, Manticoceras lamed, M. aff.lamed, Playfordites cf. tripartitus, Linguatornoceras aff. lin-guum, Aulatornoceras paucistriatum (d’Archiac & deVerneuil, 1842), and Crassotornoceras aff. crassum (Matern,1931a) (Text-figs. 15, 61).

Localities.–Locality numbers with asterisks have sections il-lustrated in Text-fig. 15.

The lower part of the Angola Shale, especially in thewest toward Lake Erie, is one of the most goniatite-rich unitsin New York. As has already been indicated (House, 1968;Kirchgasser, 1973), microcyclothemic units can be corre-lated quite widely (Text-fig. 15). Accordingly, the horizonsuffixes given to the locality numbers for this formation aregiven serially upward from the base (1-9), using the standardsequence established in Hampton Brook (Loc. 74) to thelevel of the 0.6 m (2 ft) black shale, 18.6 m (61 ft) above theblack shale immediately above the Scraggy Bed. For higherunits, and where convincing correlation is not established,the usual procedure is adopted.

Locality 71*, Farnham Creek (Lake Erie Shore).–Ap-proximately 500 yd (460 m) south of the mouth of Farnham(or Muddy) Creek (Erie County), the Angola Shale reap-pears after a concealed interval. Bed 71/9a is a horizon withpyritized goniatites (includingManticoceras lamed and Aula-tornoceras aff. eifliense) in a 0.6 m (2 ft)-thick band of blackand gray shales at the foot of the exposure. Bed 71/9b is aline of flat concretions 0.38 m (1.25 ft) higher.

Locality 72*, Point Breeze (Lake Erie Shore).–On theshore of Lake Erie, at the southern end of Grandview Bay,3.5 mi (5.6 km) south-southwest of Angola at Point Breeze(Erie County), beds low in the Angola Shale crop out nearthe point. Lake level at the northern side of the point fallswithin the lower part of the second microcyclothem abovethe black shale succeeding the Scraggy Bed, but usually theselower beds are not seen. The first horizon of large concre-tions comprises Bed 72/2, and the concretionary level in thethird microcyclothem comprises Bed 72/3, and this is ap-proximately at water level on the southern side of PointBreeze. The fauna of the Point Breeze Goniatite Bed (House,1968: 1066), withManticoceras lamed and Sphaeromantico-ceras rhynchostomum, was collected from 120 m (130 yd)south of Point Breeze (Loc. 72/6) from the top of a smallgully showing an anticlinal fold in the sixth microcyclothem.To the south, higher beds are concealed, but the thin tur-bidite in the seventh microcyclothem crops out to the southat the mouth of Farnham (or Muddy) Creek.

Locality 73*, Big Sister Creek, Angola.–The classic areafor the collection of Angola Shale goniatites by J. M. Clarkewas given solely as Angola (Erie County), but in his Guide

(Clarke, 1899c), he gave more precise localities that indicatethat the sections from which he collected are those alongBig Sister Creek on the northeastern side of Angola (ErieCounty). Many goniatite-bearing horizons are present in themicrocyclothemic succession (Text-fig. 15) and it is impos-sible to say from exactly which the type material came.

The Scraggy Bed is exposed along Big Sister Creek ap-proximately 90 yd (82 m) downstream from Mill Roadbridge, Angola. The 0.46 m (1.5 ft)-thick marker blackshale, here taken as the base of the Angola Shale, crops outapproximately 20 yd (18 m) toward the bridge. Goniatitesoccur in the concretions of the first microcyclothem. Bed73/2 is in the second microcyclothem and this is the firstline of large concretions below the Mill Road bridge. Beds73/2-4 are exposed upstream from the old mill dam to therailroad crossing and are on the western side. The 60 cm (2ft)-thick black shale is exposed in the cliff above. This seemsto be one of Clarke’s localities because he referred to sectionsbelow the railroad culvert.

The higher succession is exposed upstream of the rail-road bridge, especially on the eastern side. The 60 cm (2 ft)-thick black shale is at river level, 183 m (200 yd) upstreamfrom the railroad bridge. In the embankment above there isa 76 cm (2.5 ft)-thick black shale at the base of the eighthmicrocyclothem at a level that projects to the level of thetop of the lower arch of the bridge; 1.5 m (5 ft) below theblack shale is the same turbiditic siltstone as that exposed atthe mouth of Farnham Creek (Loc. 72). The Point BreezeGoniatite Bed (Bed 73/6) with Sphaeromanticoceras oxy, S.rhynchostomum, and Carinoceras vagans, is in the microcy-clothem (number six) immediately below the siltstone bed.A horizon of pyritic goniatites (Bed 73/9) withManticoceraslamed was located 1 ft (0.3 m) above a 20 cm (8 in)-thicksiltstone unit.

Locality 74*, Hampton Brook.–An excellent section isexposed along Hampton Brook, south of Hamburg (ErieCounty; Colton, 1956; Pepper et al., 1956: section 2). Theblack shale marker above the Scraggy Bed is at approxi-mately 760 ft (232 m) altitude, downstream of the Ham-burg-East Eden Road bridge, 1.1 mi (1.8 km) southeast ofthe confluence of Hampton Brook and Eighteenmile Creek.The first falls below the bridge are over a siltstone above a 60cm (2 ft)-thick black shale succeeding the ninth microcy-clothem. Prominent markers to lower microcyclothems arein the overhanging turbiditic siltstone with good bottomstructures in the seventh microcyclothem and a higher 0.15m (6 in)-thick siltstone in the eighth microcyclothem. Therichest faunas come from the Point Breeze Goniatite Bed(Bed 74/6b), and especially a rather lower concretionarylevel in the sixth microcyclothem (Bed 74/6a), which con-tains Sphaeromanticoceras oxy, Carinoceras vagans, S. rhyn-


chostomum, Aulatornoceras paucistriatum, and Crassotorno-ceras aff. crassum. Downstream a footpath crosses the creekon a 15 cm (6 in)-thick black shale and the concretions ofBed 74/3 in the underlying third microcyclothem yield C.vagans, A. auris (Quenstedt, 1846) group, and A. paucistria-tum.

Above the bridge the section continues up to the NorthBoston-Kehe Road bridge and beyond. “The Trinity” blackshales are exposed where a side creek joins Hampton Brookat approximately 840 ft (256 m) altitude.

Locality 75*, Cazenovia Creek.–In the West Branch ofCazenovia Creek, where the Jewettville road crosses atGriffins Mills (Erie County), a siltstone in the eighth mi-crocyclothem forms a falls beneath the bridge. There is oneprominent thin siltstone 2.5 m (8.25 ft) below which is theturbidite of Farnham Creek mouth (Loc. 72). The fauna ofthe Point Breeze Goniaite Bed (Bed 75/6) comes from aconcretionary level below the 30 cm (1 ft)-thick black shalebeneath this unit; the fauna includes Sphaeromanticocerasoxy, Carinoceras vagans, S. rhynchostomum, Linguatornocerasaff. linguum, Aulatornoceras paucistriatum, Crassotornocerasaff. crassum, and Tornoceras cf. typum. Bed 75/4 with C. va-gans and Manticoceras lamed is the upper level of large flatconcretions lower down in the fourth microcyclothem.

Locality 77*, Glade Creek.–Upstream of a bridge on theDutch Hollow Road, on Glade Creek, 0.7 mi (1.1 km)north of Strykersville (Wyoming County), there are fallsover a sandstone, 0.8 m (2.7 ft) in thickness, at an altitudeof approximately 1,075 ft (327 m), which has been taken asthe base of the Nunda Sandstone. This is marked at the topof the section illustrated here (Text-fig. 15), which was meas-ured downstream successively below the new and oldbridges. A 30 cm (1 ft)-thick black shale marks the base of“The Trinity” black shales, and Bed 77/3 is a course of con-cretions 0.48 m (1.6 ft) above it. There are two levels ofKnollenkalk siltstone below, and these comprise Beds 77/2and 77/1, the latter with Carinoceras aff. vagans. This sectionis wholly above NY Rte. 78.

Locality 78*, Sheldon Creek.–Sheldon Creek, a tributaryto Buffalo Creek, crosses NY Rte. 78, 1.8 mi (2.9 km)north-northwest of Strykersville (Wyoming County). Thetop beds of the Rhinestreet Shale are exposed above glacialdebris above NY Rte. 78 and below gravel quarries. Themarker black shale, here taken as the base of the AngolaShale, is exposed at approximately 995 ft (303 m) altitude.The best faunas are from two levels in the fourth microcy-clothem (Beds 78/4a and 78/4b) below a 61 cm (2 ft)-thickblack shale. The fauna of Bed 78/4b includes Manticoceraslamed, Aulatornoceras auris group, A. aff. eifliense, and A.paucistriatum. The Point Breeze Goniatite Bed (Bed 78/6)was located higher in the section. The creek continues the

section upward to Johnson’s Falls below the Strykersville toDutch Hollow Road.

Locality 79*, Varysburg.–The section given here is acomposite of exposures along tributaries of TonawandaCreek near the village of Varysburg (Wyoming County)(Text-fig. 15; Locality 80 read 79). The lowest beds (Beds79/1-4) are exposed in the floor and at the edge ofTonawanda Creek, downstream of the US Rte. 20A crossingon the western side of the village. Bed 79/4 is characterizedby occurrences ofManticoceras lamed. The section continuesupward to levels near the Point Breeze Goniatite Bed (Bed79/6) in the gully of the tributary west of the village belowa disused railroad track; Clarke (1899c: 110) reportedMan-tioceras (= Sphaeromanticoceras) oxy from approximately thelevel of the tracks. The upper part of the illustrated section(Beds 79/11-13) is taken from exposures upstream of therailroad track in which “The Trinity” black shales can bereadily located beneath a 0.66 m (2.2 ft)-thick siltstone atthe top of a falls. Massive siltstones typical of the NundaSandstone enter 7 m (23 ft) above the top of the falls, andthe Pipe Creek Shale (and top of Angola) projects to a levelapproximately 41.5 m (136 ft) higher. On the section ofPepper et al. (1956: section 5), only the upper parts of thissection were referred to the Nunda, and the lowest siltstoneswere marked as West Hill Member. It should be noted thatthe marker black shale that defines the base of the AngolaShale (and top of Rhinestreet Shale) is probably one of thetwo lowest black shales marked by Pepper et al. (1956) intheir section 5.

The middle part of the illustrated section, from the PointBreeze Goniatite Bed (Bed 79/6) upward to Beds 79/9 and79/10, a Knollenkalk horizon with Carinoceras aff. vagans,Manticoceras lamed, and “The Trinity” black shales, is basedon exposures along Stony Brook east of Varysburg, the eas-iest access to which is a farm track that crosses the middle ofthe section. Part of the upper section was measured in a sidebranch that joins Stony Brook at approximately 1,225 ft(374 m) altitude.

Locality 80, Varysburg (Gassman Road).–Within theNunda interval above the section measured for Text-fig. 15is a Knollenkalk horizon that has yielded a single Sphaero-manticoceras rhynchostomum?. The horizon (Bed 80/14), atapproximately 1,375 ft (419 m) altitude, is 1.8 m (6 ft)above the old bridge foundation where the Gassman Roadcrossed Stony Brook east of Varysburg (Wyoming County).The horizon, within highly burrow shales with Zoophycostraces, might be the middle or upper of the three nodulebands in the Nunda Sandstone interval shown in section 5of Pepper et al. (1956).

Locality 81*, Johnson Creek.–Approximately 2.4 mi (4km) north of Varysburg, on Johnson Creek (Wyoming


County), a long section of Angola equivalents continuesabove Loc. 57. The marker black shale of the basal Angolais in the middle of a cascade approximately 350 yd (320 m)west of disused railroad west-northwest of Earls. The PointBreeze Goniatite Bed (Bed 81/6), with Sphaeromanticocerasrhynchostomum and Crassotornoceras aff. crassum, comprises

concretions near the top of the sixth microcyclothem, ap-proximately 3.6 m (12 ft) above the top of the falls over silt-stones in the fourth microcyclothem. Bed 81/10 can berecognized by a 0.25 m (10 in)-thick mudrock unit withtrace fossils that forms a ledge across the creek, 120 yd (110m) below the north-south Maxon (Beck) Road on the hill

Text-fig. 16. Sections of the Hanover Shale equivalents between Walnut Creek at Silver Creek (Chautauqua County) and Beaver MeadowCreek at Java Village (Wyoming County), with the locality numbers used in the text. Note: The 20 cm concretionary bed below the F/Fboundary in Irish Gulf is Loc. 90/2, and the black shale in the lower Hanover in Glade Creek (Loc. 91) is 84 (not 64) cm thick.



above.Locality 82*, Relyea Creek.–The section in Relyea Creek

(Text-fig. 15) continues the Rhinestreet sequence of Loc. 60(Text-fig. 13) and commences 0.6 m (2 ft) above the base ofa main waterfall, 650 yd (595 m) west of NY Rte. 19, atSouth Warsaw (Wyoming County). Bed 82/6, the PointBreeze Goniatite Bed, is approximately 7.6 m (25 ft) abovethe base of the marker black shale at the base of the Angolaand approximately 8.5 m (28 ft) below the top of the mainwaterfall; the fauna includes Sphaeromanticoceras rhynchosto-mum,Manticoceras aff. lamed, Playfordites cf. tripartitus, andCrassotornoceras aff. crassum.

Locality 83*, Stony Creek.–There is a fine section inStony Creek (Text-fig. 15) and the section given here con-tinues upward from Loc. 61 (Text-fig. 13). The basal markerblack shale of the Angola is seen in the face of the main wa-terfall below the railroad bridge 2,200 yd (2 km) southwestofWarsaw (Wyoming County). The 15 cm (6 in)-thick silt-stone on which the railroad bridge rests is a convenientmarker for Bed 83/6a, which has yielded a very fine faunawith Manticoceras aff. lamed and Sphaeromanticoceras rhyn-chostomum. Another bed, 83/6b, is 0.7 m (2.3 ft) above thissiltstone. Bed 83/8 is a silty Knollenkalk horizon withSphaeromanticoceras oxy immediately above a 18 cm (7 in)-thick siltstone, 3.6 m (12 ft) above the bridge foundationunit.

Locality 84*, Kennedy Gulf.–Loc. 84 is a continuationinto the lower Angola Shale of the section in the UpperRhinestreet at Kennedy Gulf (Loc. 63; Text-fig. 13), southof Dale (Wyoming County). A concretion in Bed 63/4yielded a fine specimen of Spaeromanticoceras oxy collectedby J. Kralick (NYSM 16536). Bed 63/4 is 2.23 m (7.3 ft)above the base of the Angola Shale (Text-fig. 13).

Locality 85, Wolf Creek.–There are falls along WolfCreek over the Nunda Sandstone just below the village ofCastile (Wyoming County). A locality approximately 250yd (230 m) downstream of the falls yielded a goniatite faunafrom grey silty mudrock that includesManticoceras lamed. Anumber of the large specimens (?Sphaeromanticoceras) haveauloporid encrustations similar to those seen on goniatites atVarysburg in levels just above “The Trinity” black shales(Bed 79/12). We were not able to demonstrate that it is thesame horizon, but estimates based on the regional dip sug-gest it is a possibility.

Locality 86*, Genesee Gorge.–The horizons at which go-niatites have been located in this survey in the GeneseeGorge (Wyoming and Livingston counties) are illustratedin Text-fig. 12. In this section, the well-known Table RockSandstone at the Lower Portage Falls is visible. The unit wasnamed by Chadwick (1933: 96) but it was first recognizedby Hall (1843: 391). In the column given by Pepper et al.

(1956: sections 11-12), this level was not indicated, but oursurvey indicates that it is the sandstone they marked approx-imately 73 m (24 ft) below the top of their section 11, andapproximately 15 m (50 ft) above the bottom of their sec-tion 12. It lies at an altitude of approximately 910 ft (277m).

The level from which the syntype of Sphaeromanticocerasoxy (NYSM 3746) came is only approximately determined,as has been indicated above, but would appear to be fromthe area of the Lower Portage Falls at the section indicated.This probably lies within late Rhinestreet Shale equivalents.

Locality 87, Stony Brook Glen.–The long section inStony Brook Glen, through the state park in SteubenCounty, south of Dansville (Livingston County), was fig-ured by Pepper et al. (1956: section 18). This is a locality ofClarke (1898: 78) for Sphaeromanticoceras oxy, but the pre-cise level was not recorded by him. There is a level of largeconcretions in siltstones with thin fossiliferous lenticles justbelow and downstream of the old railroad bridge, but goni-atites were not noted there.

Locality 87a. I-390–Dansville.–The Dansville area(Text-fig. 3) is also the source of a specimen of Sphaeroman-ticoceras oxy (NYSM 16570) from the upper West FallsGroup (Nunda or Wiscoy Sandstone equivalents) collectedfrom loose blocks by B. Oldfield (Broome Community Col-lege). The horizon is a shell-rich channel fill on the southernside of US Rte. I-390 (between markers 15 and 14) nearExit 4 (Wayland), in Steuben County, south of Dansville(Livingston County). Stop A-5-4, New York State Geolog-ical Association (Kirchgasser, Over, &Woodrow, 1994: 358)and Stop B-9, Subcommission on Devonian Stratigraphy(SDS) (Kirchgasser, Over, & Woodrow, 1997: 232).

Pipe Creek Shale and Hanover ShaleThe black Pipe Creek Shale and gray Hanover Shale werecombined as the Java Formation by de Witt (1960) but weare following older usage and including these units as part ofthe upperWest Falls Group. Details of the lithostratigraphicvariation in these units were worked out by Pepper & deWitt (1950) and considerable detail was given on theirchart. Correlations within the Pipe Creek, Hanover, andlowermost black Dunkirk shales of the succeeding Canad-away Group in Erie and Wyoming counties have been re-fined by Over (1997a, b) and Over et al. (1997). A detailedtransect of uppermost Hanover and lowermost Dunkirk cor-relations between Lake Erie and the town of Java inWyoming County was also provided by Baird & Lash(1990: fig. 5).

The Pipe Creek Shale forms a clearly marked basal unitof black shale that is only 0.6 m (2 ft) thick on WalnutCreek (Locs 89, 89a; Text-fig. 16), south of the village of

Silver Creek, near Lake Erie in Chautauqua County, but theunit thickens eastward and Sutton (1963) recognized thehorizon as far east as the Waverly and Owego Quadrangles(Text-fig. 1). No goniatites have been reported from the PipeCreek Shale and none were noted in this study.

The Hanover Shale, a medium-gray shale with blackshale bands and siltstones especially in the upper part, is 29m (94 ft) thick on Walnut Creek (Loc. 89a), a locality thathas produced the best goniatites faunas in the lower part ofthe unit, mostly small Dephiceras cataphractum (Clarke,1898) (Beds 89a/2-4) but including Crickites lindneri(Glenister, 1958) from a level located by G. Kloc in thevicinity of Beds 89a/4 or 89a/5 (Text-fig. 16). Other loosespecimens of C. lindneri collected by G. Klapper (NYSM16537 and 16538) and R. T. Becker (HM MBC 1943)probably came from the same level. Bed 89a/5 or just aboveis the probable source of the specimen collected by G. Klocthat we earlier described as Sphaeromanticoceras aff. rickardi(House & Kirchgasser, 1993: text-fig. 56G-I). G. Kloc hasa collection of pyritic goniatites from the Pipe Creek-Hanover contact beneath the NY Rte. 20 bridge in WalnutCreek (Loc. 89a), but the collection has yet to be described;this pyritic level was apparently covered by creek sedimentwhen we did our survey. The same levels near the PipeCreek-Hanover contact outcropping in Silver Creek, up-stream of the NY Rte. 20/5 bridge in the village of SilverCreek, have yielded goniatites to collectors but this localitywas not included in our survey. The lowermost Hanover isthe source of the cotypes of Delphiceras cataphractum andprobably also of the lectotype (NYSM 4092) and syntypesof Aulatornoceras rhysum (Clarke, 1898), which came fromBeaver Meadow Creek (Loc. 92), at Java in WyomingCounty.

At a higher level in the Hanover Shale at Glade Creek(Loc. 91) near Strykersville in Wyoming County, L. V.Rickard found a specimen of Sphaeromanticoceras rickardi(NYSM 12032) from a concretionary bed (Bed 91/7) withina series of silty beds. From near the top of the Hanover Shalein Irish Gulf (Loc. 90), ?Archoceras sp. (NYSM 16587 and16588) and ?Crickites sp. juv. (NYSM 16589) occur in Bed90/2, a level that also yields conodonts of MN Zone 13(Over, 1997a, b). These are the highest known goniatites inthe Frasnian succession in New York State.

Eastward the Hanover Shale passes progressively into theWiscoy Sandstone (Pepper & deWitt, 1950) and atWiscoyCreek (Loc. 93) goniatites have been reported but none werefound in this study. Luther (1911: 23) stated thatMantico-ceras (= Sphaeromanticoceras) oxy occasionally appears in theWiscoy beds.

The precise relations of the Frasnian/Famennian bound-ary have been elucidated by J. Over at several localities using

conodonts, particularly in the sections in Irish Gulf (Loc.90) and Beaver Meadow Creek (Loc. 92) and the interpre-tations shown onText-fig. 16 follow his work (Over, 1997a,b; Over et al., 1997).

Localities.–Locality numbers with asterisks have sections il-lusrated in Text-fig. 16. Horizons bearing goniatites in theHanover Shale are restricted to the western outcrops close toLake Erie and only a few sections were measured by us(Text-fig. 16).

Locality 88, Eden Valley.–The South Branch of Eigh-teenmile Creek passes through Eden Valley (Erie County);2.5 mi (4 km) upstream of Eden Valley where the ChurchRoad bridge crosses the creek. Approximately 200 yd (183m) upstream is the old bridge where the former road crossedbeside a sawmill at approximately 800 ft (244 m) altitude.The Pipe Creek Black Shale lies below. The Pipe Creek ishere approximately 1.2 m (4 ft) thick, and the overlyingbasal Hanover Shale has small concretionary nodules withmanticoceratids. Immediately above follow thick black-shaleunits.

Locality 89a*, Walnut Creek.–On Walnut Creek southof the village of Silver Creek (Chautauqua County), there isa long and fine section of the Hanover Shale. The black PipeCreek Shale crops out below the NY Rte. 20 bridge and thesection figured is upstream of the bridge where the basalbeds of the Hanover Shale are well exposed for several hun-dred yards (meters) on the eastern side of the creek. Bed89a/4, the most productive horizon, is at stream level 40 yd(37 m) above the bridge and has a thick black shale, 0.3 m(1 ft) above it. Beds 89a/1-3 are best exposed approximately400 yd (365 m) farther upstream. Beds 89a/2-4 containDel-phiceras cataphractum and Bed 89a/5 yielded to G. Kloc afine specimen of Sphaeromanticoceras aff. rickardi and oneof Crickites lindneri. Bed 89a/5 is probably the source of thetwo loose specimens of C. lindneri described herein.

Locality 89*, Walnut Creek.–Higher upstream on Wal-nut Creek in Chautauqua County, above the concealed in-terval marked by Pepper & de Witt (1950: section 1), theupper beds of the Hanover Shale are exposed. No goniatiteswere collected from these levels and the placing of the Frasn-ian/Famennian boundary is based on that by Over (1997b).

Locality 90*, Irish Gulf.–A fine section of Hanover Shalecommences in Irish Gulf upstream of the railroad crossingabove where NY Rte. 391 (Boston State Road and old NYRte. 219) crosses the creek, 0.3 mi (0.5 km) south of theChestnut Ridge Road and 1.6 mi (2.6 km) southeast ofNorth Boston (Erie County; Text-fig. 16). Goniatites withmanticoceratid sutures occur in the concretions (with whitebarite) of Bed 90/1 (WTK 4100) in the creek floor at thelevel of the gas seeps, approximately 2.5 m above the base of


the Hanover; conveivably these could be large specimens ofDelphiceras cataphractum but the inner whorls are not pre-served. The highest known goniatites in the New YorkFrasnian occur in Bed 90/2, a 2.0-2.5 cm-thick concre-tionary horizon capping a small falls in the upper HanoverShale, 4.7 m below the base of the conodont-defined Frasn-ian/Famennian boundary (Over 1997a, b). They consist ofsmall, poorly preserved whorl sections of ?Archoceras sp.(NYSM 16587, 16588) and ?Crickites sp. juv. (NYSM16589). This baritic crinoid and gastropod-rich horizon wasillustrated by a concretion symbol by Over (1997b: fig. 8).

Locality 91*, Glade Creek.–North of Strykersville, inGlade Creek (Wyoming County), the middle and upperparts of the Hanover Shale are well exposed between theStykersville Road north to Dutch Corner and the PlantsCorner Road to Sheldon. The basal black Pipe Creek Shalecrosses the creek at approximately 1,220 ft (372 m) altitude,and the basal black Dunkirk Shale at approximately 1,350ft (411 m). In the lower part of the section a boundary fencecrosses the creek and a short distance above is a 0.84 m (2.75ft)-thick black shale marker band with a thin siltstone 2 m(6.5 ft) higher. A loose specimen of Sphaeromanticocerasrickardi (NYSM 12033) was found in this part of the lowerHanover Shale. Bed 91/1 is a horizon of small nodules be-tween two thin black shales, 0.69 m (2.25 ft) apart, whichlie above the siltstone. At 0.6 m (2 ft) above the higher thinblack shale is a broad spread of small concretions forming aledge across the stream, which constitute Bed 91/2. Bed91/4 is at the foot of the falls above. Bed 91/7, a 10-15 cm(4-6 in)-thick Knollenkalk siltstone that produced a singlespecimen of S. rickardi (NYSM 12032), forms the crest ofa 0.6 m (2 ft)-high waterfall, 1.6 m (5.25 ft) above a large4.5 m (15 ft)-high waterfall. Bed 91/7 is approximately 7.6m (25 ft) below the base of the Dunkirk Shale.

A track down from the road between Plants Corner andSheldon gives the easiest access to the upper part of the sec-tion. The track crosses the creek almost at the Hanover/Dunkirk contact with a 15.2 cm (6 in)-thick hard limestoneband 0.6 m (2 ft) below. The Huddle Rider Bed is a thinblack shale with possible cephalopod anaptychi, 0.76 m (2.5ft) below the limestone band.

Locality 92*, Beaver Meadow Creek.–Above Angel Falls,in the town of Java (Wyoming County), a long and inter-mittent section is exposed along a tributary to Buffalo Creek(Pepper & deWitt, 1950: section 6). NY Rte. 78 crosses thecreek just below Angel Falls (over the Nunda Sandstone)and approximately 150 yd (137 m) above the road the upperpart of the Pipe Creek Black Shale forms a small falls. Bed92/1 is immediately above the falls on the northern bank.On the opposite side of the creek there is a ford crossing toa small island and Bed 92/2 is on the island side of the ford.

Bed 92/2 is probably the type horizon of Delphiceras cat-aphractum and Aulatornoceras rhysum. A bed of larger con-cretions, 91/3, crosses the creek upstream, below the remainsof a wooden dam.There is a considerable section exposed onthe northern bank of the creek. Bed 92/4 is a horizon oflarge concretions, 0.9 m (3 ft) above stream level at the footof this section.

Locality 93,Wiscoy Creek.–Equivalents of the HanoverShale tongue into the Wiscoy Sandstone along WiscoyCreek,Wiscoy (Allegany County), below the dam. A visit bythe Friends of the Devonian in 1962 (J. W. Wells, pers.comm.) found goniatites near the second cascade, but nonewere located by us.

CANADAWAY GROUP

The Canadaway Group (Chadwick, 1933) comprises thenext sequence initiated by the entry of another prominentblack shale, the Dunkirk Shale, which is succeeded by theSouth Wales Shale, Gowanda Shale, Laona Sandstone, anda series of other units that have not yielded goniatites and donot concern us here. Pepper & deWitt (1951) provided de-tailed sequences through the group and Baird & Lash(1990), Over (1997a, b), and Over et al. (1997) added sig-nificant refinements, particularly in the lower DunkirkShale. Goniatites have only been discovered in the GowandaShale during this work at two levels, a level with Truyolsocerascf. bicostatum (Hall, 1843) (Loc. 99) on Walnut Creek anda rather higher level that has been termed the Corell’s PointGoniatite Bed (Loc. 97) with Cheiloceras (Cheiloceras) ambly-lobum (G. & F. Sandberger, 1851), Phoenixites concentricus(House, 1965), and Truyolsoceras bicostatum. The level nearForestville which yielded T. clarkei (Miller, 1938) has notbeen located. When the entry of Cheiloceras in Europe wastaken as a marker to the lowest Famennian, the recognitionof this level in New York (House, 1962) provided the firstconvincing evidence for the location of the Frasnian/Fa-mennian boundary, but a Global Stratigraphic Section andPoint has now been established by the International Unionof Geological Sciences (IUGS) for the boundary in theMontagne Noire, France (Klapper et al., 1993; House,Becker et al., 2000) at a significantly older level. In NewYork, the boundary is best recognized using conodonts(Over, 1997a, b), as commented upon earlier (Text-fig. 16).Gephuroceratids are last seen just below the boundary inIrish Gulf (Loc. 90), as recounted earlier, at approximatelythe level where the Gephurocerataceae become extinct in-ternationally.

The Corell’s Point Goniatite Bed has been located at anumber of localities (House, 1968) and these are indicatedon Text-fig. 17. It is clear that the locality at Corell’s Point(Loc. 97) was known to James Hall and this could be the


source of the type of Truyolsoceras bicostatum that he de-scribed in 1843. Cheiloceras is present in collections made inthe last century and held in the NYSM but the specimens re-mained unrecognized (House, 1962: 276). Despite substan-tial collecting at the new localities. no additional goniatitespecies have been added as a result of our current work.

Localities.–Locality numbers with asterisks have sections il-lustrated in Text-fig. 17. The Corell’s Point Goniatite Bed(House, 1968: 1066) is almost the only horizon in theGowanda Shale known to carry goniatites, but there is arather lower level known in Walnut Creek (Loc. 99). TheCorell’s Point Goniatite Bed has been traced from the typelocality on Lake Erie Shore (Chautauqua County) eastwardto Gowanda (Erie-Cattaraugus counties), and faunal andlithological evidence for the horizon has been found as fareast as Java (Wyoming County). Tesmer (1954, 1975) gaveinformation on Gowanda localities in Chautauqua and Cat-taraugus counties. The main goniatite localities are as follows(Text-fig. 17):

Locality 97*, Corell’s Point.–Corell’s Point lies between

the outlets of Walker and Corell’s creeks, 250 yd (230 m)southwest of the former, 2.85 mi (4.6 km) west of Brocton(Chatauqua County) at an altitude of approximately 572 ft(174 m). A small point there is formed by a ledge of septar-ian nodules, the Corell’s Point Goniatite Bed, to 0.46 m (18in) in thickness, overlain by 4.6 m (15 ft) of shale with cal-careous concretions. The fauna occurs in the ledge and as-sociated shale. The goniatites include Cheiloceras (C.)amblylobum, Truyolsoceras bicostatum, and Phoenixites con-centricus, the last a species described by House (1962, 1965)under Tornoceras, but which is now referred to Phoenixites(Becker, 1993a).

Although apparently known to James Hall (1843: 246),and probably the source of his Goniatites bicostatus Hall,1843, the first clear reference to the bed seems to be byLuther (1903: 1025) who referred to concretions on theshore west of Brocton that could refer to this locality. TheClarke and Luther collections of Cheiloceras (C.) amblylobumdated 1898 are at Albany (NYSM 11239-11242). The redis-covery of this locality in recent times is due to W. Moranwhose collection is in Washington, DC (USNM 137665

Text-fig. 17. Sections of the localities at which the fauna of the Corell's Point Goniatite Bed in the Gowanda Shale has been located betweenLake Erie and Java Village (Wyoming County), with the locality numbers used in the text.



and 137666). This section was also illustrated by Baird &Lash (1990: fig. 4).

Locality 98*, Little Canadaway Creek.–Eastward theCorell’s Point Goniatite Bed crosses Little CanadawayCreek, 2,200 yd (2 km), north-northwest of the junction ofLake Road and NY Rte. 20 at Lamberton (ChautauquaCounty) at an elevation of approximately 625 ft (190 m).The section lies above the creek crossing of the New YorkState Thruway. A bioturbated 12.7 cm (5 in)-thick siltstonebed is seen approximately just where exposures commenceabove the thruway overpass, and the goniatite bed is approx-imately 3.9 m (13 ft) higher. The calcareous ledge is againseptarian and of irregular thickness, to 0.38 m (15 in). Thesame fauna recurs as at Corell’s Point.

At the mouth of Little Canadaway Creek, 1.4 mi (2.3km) south-southwest of Van Buren Point on Lake ErieShore, blocks of the Corell’s Point Goniatite Bed with thetypical fauna can be found.

Locality 99*, Walnut Creek.–OnWalnut Creek, there isa long section of the Gowanda Shale (Pepper & de Witt,1951: section 1). The first horizon bearing goniatites wasfound at a level 0.9 m (3 ft) above the siltstone on whichthe Mixer (Sheridan) Road bridge (Loc. 99/a) is built, onthe northern outskirts of Forestville (Chautauqua County).This level yields crushed Truyolsoceras cf. bicostatum moreevolute than the type described by Miller (1938: 80) asTornoceras (Aulatornoceras) clarkei. The locality of Truyol-soceras? clarkei was given by Miller as near Forestville, NewYork, but it could be that the material he described was thatreferred to by Harris (1899) as from Walnut Creek, 9.1 m(30 ft) below the railroad bridge and approximately 50 m(10 rods or 165 ft) downcreek from it. This is in the upperpart of the Gowanda, and not the Laona Sandstone orWest-field Shale, as speculated earlier (House, 1965: 123), but thematerial might not be the same.

The Corell’s Point Goniatite Beds, exposed in WalnutCreek approximately 300 yd (275 m) upstream from therailroad bridge at Forestville (Loc. 99b), yields Truyolsocerasbicostatum and Cheiloceras amblylobum. The elevation of thebed is approximately 847 ft (258 m). This locality also seemsto have been noted by Harris (1899), who recorded fossilsassociated with concretions approximately 100 m (20 rods)above the railroad culvert. On section 1 of Pepper & deWitt(1951), the goniatite bed is marked as a layer of concretions1.5 m (5 ft) above a 0.6 m (2 ft)-thick black shale, 18 m (59ft) below the base of the Laona Sandstone.

Locality 100, Smith Mills.–In Chautauqua County, theCorell’s Point Goniatite Bed is exposed again approximately150 yd (137 m) upstream from the King Road bridge cross-ing of a tributary of Silver Creek, approximately 0.25 mi(0.4 km) west of Smith Mills at an altitude of approximately

865 ft (264 m). The sequence of the Gowanda Shale cannotbe measured in detail here, but the distinctive fauna andlithology of the goniatite bed make the assignment clear.

Locality 101, Big Indian Creek.–On Big Indian Creek(Cattaraugus County), the Corell’s Point Goniatite Bedcrops out 46 m (50 yds) below a track and ford crossingfrom Olmstead Farm, approximately halfway betweenWardtown Road bridge and Planck (or Plank) Road at an al-titude of approximately 880 ft (268 m).

Locality 102, Little Indian Creek–The locality on LittleIndian Creek (Cattaraugus County) lies approximately 64 m(70 yd) downstream from the crossing of Planck Road, at analtitude of 890 ft (271 m). Loose blocks of the unit are com-mon for 30 yd (27 m) farther downstream, from which thegoniatites mentioned here were collected.

Locality 103, Cattaragus Creek Tributary.–On the Cat-taraugus Indian Reservation, the Corell’s Point GoniatiteBed was located just above the Mackinaw Road Bridgecrossing of a tributary of Cattaraugus Creek, 3 mi (4.8 km)northwest of Gowanda (Cattaraugus County) at an altitudeof approximately 900 ft (274 m).

Locality 104, Cattaragus Creek, South Branch.–Thereis a long and excellent section of the Gowanda Shale alongthe South Branch of Cattaraugus Creek, southeast ofGowanda (Cattaraugus County). The Corell’s Point Goni-atite Bed forms a joint-controlled water-shoot (or flume), 1mi (1.6 km) south-southeast of Forty Bridge, 675 m (739yd) south along the creek from a stream and waterfallmarked on the Gowanda 7 1/2’ Quadrangle Map, at an al-titude of 975 ft (297 m). A good goniatite fauna was col-lected from this locality.

Locality 105, Clear Creek, North Branch, Taylor Hol-low.–Approximately 2.5 mi (4 km) east of Taylor Hollow(Cattauraugus County) is a tributary of the North Branch ofClear Creek. Approximately 150 yd (140 m) above theLenox Road, blocks of the Corell’s Point Goniatite Bed with?Cheiloceras were located at approximately 970 ft (296 m) al-titude. The associated beds here are siltier than those fartherwest.

Locality 106, Clear Creek, North Branch,Marshfield.–In Erie County, 2 mi (3.2 km) west-southwestof Marshfield, on the North Branch of Clear Creek, 100 yd(90 m) downstream from Jenings Road bridge, a soft silt-stone immediately above a concretionary layer containsTruyolsoceras cf. bicostatum in a setting similar to that east ofTaylor Hollow. The altitude is approximately 1,050 ft (320m).

Locality 107, Anthony Gulf.–On the eastern side of An-thony Gulf, 2.4 mi (4 km) north-northeast of Boston Cen-ter (Erie County), Cheiloceras was found in the typicalrecrystallized preservation of the Corell’s Point Goniatite

Bed, just below a pipeline crossing at an altitude of 1,350 ft(411 m). The block was not in situ, but can scarcely havebeen far displaced.

Locality 108, Gears Creek.–On Gears Creek, 1.8 mi (2.9km) south-southwest of Holland (Erie County), a looseblock of the Corell’s Point Goniatite Bed with ?Phoenixitessp. in the usual preservation was found at an altitude of ap-proximately 1,400 ft (427 m).

Locality 109, Java.–A specimen of Cheiloceras (Cheilo-ceras) amblylobum in the NYSM is labeled “Portage, Java,”presumably referring to a locality near Java (WyomingCounty). Despite a careful search, the Corell’s Point Goni-atite Bed was not located in place. H. S. Williams (1887:49), however, gave a section in which he recorded goniatitesand this deserves further attention. Java is the farthest eastthat Cheiloceras has been identified. Two specimens ofPhoenexites concentricus (NYSM 11965 and 11966) arethought to have come from Gowanda Shale equivalents atJava.

CONNEAUT GROUP

Only two goniatites are recorded from the Conneaut Groupof New York State. The only specimen of these that can besatisfactorily determined is USNM 137645, described hereas Maeneceras aff. acutolaterale (G. & F. Sandberger, 1850)and formerly assigned to Maeneceras cf. pompeckji(Wedekind, 1918) (House & Kirchgasser, 1993: 276). Thelocality is in the Ellicott Shale at Porter Creek (Loc. 110),near Summerdale (Chautauga County). The other specimen(NYSM 4090) is the holotype of Tornoceras edwinhalliClarke (1898: 111, text-fig. 85), which is recorded as fromthe Conneaut at Nile (Allegany County). A subsequent ob-servation by G. H. Chadwick reported by Miller (1938:152) suggests that the specimen could be from either theChagrin or Volusia member of the Conneaut; the level isnot satisfactorily determined. The specimen is describedbelow as “family and genus indeterminate.”

Localities.–Locality 110, Porter Creek, Summerdale.–A locality near

Summerdale (Chautauqua County) was the source of a go-niatite collected by H. S.Williams (his locality 518B; detailsare in his notebooks preserved at the USGS, Washington,DC). The specimen (USNM 137654) is here referred toMaeneceras aff. acutolaterale. J. W. Wells found the locality518B marked on a map belonging to H. S. Williams. Thelocality is 2 mi (3.2 km) “northwest of Summerdale alongPorter Creek at a point just below Porter Cemetery and atan altitude of 1450 feet [442 m]” (House, 1962: 277). Onthe geologic map of Tesmer (1954: 29), the locality falls inthe northern (middle) part of the outcrop of the Ellicott

Shale. This is thought to be a Famennian IIB level in termsof the German sequence (House, 1962) or UD II-G in moremodern terms (Becker & House, 2000).

CONEWANGO GROUP

No goniatites are known this high in the sequence in NewYork State but faunas are known in adjacent Erie County,Pennsylvania (Loc. 111, below) and there are three Famenn-ian goniatite and clymenid faunas known near Cleveland,Ohio, in the Cleveland Shale and earliest Bedford Shale(House et al., 1986). According to Oliver et al. (1969), thebase of the Cleveland Shale where the lowest of the threeCleveland faunas occurs (just above the extreme western-most tongue of the Chagrin), lies at approximately the levelof the Wolf Creek and Panama conglomerates of New YorkState, and hence all three Cleveland faunas would be referredto the Conewango Group. No attempt has been made inthis work to trace the Ohio levels eastward into Pennsylvaniaand New York.

Localities.–Locality 111, Howard Quarries.–North-northeast of

Franklin Corners, 14 mi (22 km) southwest of Erie (ErieCounty), Pennsylvania, at Howard Quarries, is the type lo-cality ofMaeneceras milleri (Flower & Caster, 1935). White(1881: 103) described the quarries here and noted that FallsRun flows northward past the quarry. The horizon was saidto be “probably lower Conewango, near the horizon of thePanama conglomerate” (Miller, 1938: 176). The lower Ve-nango Sandstone is said to occur just below the quarry out-crops. The specimen was determined by House (1962: 262)as Sporadoceras milleri, but would now be referred toMaeneceras (following Becker, 1993a) and is so describedhere; it is correlated with the Cheiloceras Stufe of Europe(UD II-H); the level of the lowest of the three Clevelandfaunas, referred to above, is referred to the PlatyclymeniaStufe (UD IV-A).

PALEONTOLOGYThis section comprises introductory discussions on the gen-eral problems relating to this work and a review of the localstratigraphic and ecological setting of the late Devonian go-niatite faunas in New York State. A review of the evolutionof Devonian ammonoids, as it applies to the faunas underdescription, and an account of the terminology and proce-dures adopted in this work are followed by the systematicdescription of the faunas themselves. The last, which consti-tutes the major section, is intended to review all the faunasknown to us and to illustrate type material photographicallyfor the first time. Only brief descriptions and comments onnew additional information are given in the cases of those


taxa for which good accounts are already available in the lit-erature.

The prime purpose of this work was to establish the de-tailed biostratigraphic occurrence of late Devonian am-monoids in New York State. That the New York goniatitefaunas were remarkably diverse and well developed has longbeen known. However, the localities of most of the knownfaunas were in only the crudest way related to each other ina biostratigraphic sense. Hence the necessity for a detailedestablishment of the stratigraphical relations of the manysections available for study, and review of these has beengiven in the previous section. A start is made in this sectionto give a more detailed analysis of the available goniatite fau-nas, and to this end general comments on the stratigraphicoccurrence and ecology of the faunas are given. Little infor-mation of this type is available for Devonian faunas else-where in the world largely because most classical Devonianammonoids are from pelagic facies. Some evidence will beprovided suggesting that there is rather more facies controlof goniatite distribution than has generally been conceded,particularly with regard to the Triainoceratidae.

The goniatite sequence for Frasnian equivalents that isnow established in New York is one of the more detailedknown in the world, and enables a more critical look at var-ious aspects of goniatite evolution. There is considerablescope for additional work of this type even in New York,and substantially more in areas with other good sequences ofthis age, as in Belgium, Germany, North Africa, WesternAustralia, and the Timan region of Russia especially. In asense, then, the present study is a start only. There are a largenumber of late Devonian genera that have not been foundin New York or elsewhere in North America, and hence notplaced in the time sequence here established. It is also truein some cases that there are no critical grounds for placingthe missing genera into the sequence in other ways usingcorrelative methods. It is to be hoped that in the future, con-odont studies of type material and other means will enablecloser time documentation of goniatite taxa.

Future contributions will depend largely on new collect-ing. We are conscious that only limited time has been spentextending the geographical range of known levels of goni-atite occurrence. Also there are certain levels that have notbeen searched systematically, notably the black shales them-selves. But the data recorded here have been assembled over40 yr and it seems unlikely that delay in recording the datawould serve much purpose. Rather, it is hoped, this reportwill encourage others to study particular levels or problemsin New York and work toward compiling similarly detaileddata on Devonian goniatite occurrences in other areas of theworld.

PALEOENVIRONMENTS

Three sedimentological controls influence the nature of thelate Devonian rocks in upstate New York, and hence the dis-tribution of goniatites. (1) In space is the dominance of thethick clastic sequences of the Catskill Delta in the east, andassociated with it, the regular reduction in thickness, andfiner grade of sediment encountered westward where, in theearlier parts of the sequence, substantial unconformities canbe documented (Rickard, 1975; Baird, 1979; Baird & Brett,1986a, b). (2) In time is the influence of a series of transgres-sive pulses, commencing with that of the Tully Limestone,and later by black shale tongues extending eastward into theclastic facies and represented successively by the black shalesof the Geneseo, Middlesex, Rhinestreet, Pipe Creek, andDunkirk members (Text-fig. 2); the onset of four of the lastare mirrored in the lithostratigraphical nomenclature byforming the base of group units (Rickard, 1975). The cyclesthus initiated are essentially shallowing-upward sequences.(3) In general, these easterly extending transgressive pulsesshow decreasing effect through time. The first here consid-ered, that of the Upper Tully, which initiated the TaghanicOnlap, extends the farthest. The later black shale tonguesare progressively more limited to western areas. All of this isillustrated onText-fig. 3. Corresponding with this, the clas-tic facies belts extend farther to the west with time, as hasbeen recognized for many years (Rickard, 1964).

Goniatite occurences are intimately linked to certain fa-cies belts that change their position, during the time periodconsidered, in relation to the transgressive pulses. Hence itis convenient to give generalizations concerning goniatitedistribution and paleocology under headings of lithologicalfacies types. Later sections will deal with geographical distri-butions and with several distinctive organic associations.

Facies TypesThe major facies belts typical of the sequence from theTaghanic to Chautauquan are illustrated in Text-fig. 18,which is based in part on Bowen et al. (1974). The diagramis largely based on the Sonyea Group, and the geographicallimits indicated apply to that group only.

Clastic facies.–Most of the clastic facies discussed fit readilyinto the sedimentological model illustrated in Text-fig. 18.The carbonate-dominated facies, however, apart from theconcretions, generally do not, and each merits separate con-sideration in the text.

(1) Black Shale.–This facies of black to grayish-black orbrown to olive-black shale, often massive and weatheringwith extremes of fissility, characterizes the major and minorblack shale tongues between the Geneseo Shale Member andthe Dunkirk Shale Member (Text-fig. 3). The thicknesses


range from a few millimeters to the great sequence of theRhinestreet Shale Member, which reaches over 60 m (200 ft)at outcrop and is very extensive in the subsurface (Rickard,1975, 1981, 1984). Some documentation of the precise col-ors involved was given by deWitt & Colton (1978).Withinthe generally black shales, lighter bands can occur, and thesegenerally enter and develop eastward, splitting up the se-quences. As will be apparent fromText-figs 3, 8, 11, 13, 15,and 16, these black shales occur as the initiating facies inboth minor sedimentary rhythms and the large sedimentarycycles, or rhythms, which define the groups.

Goniatites are very rare in this facies with the exceptionof a few horizons. Whether this rarity is due to early diage-netic dissolution of the aragonite goniatite shells has notbeen determined. Levels with crushed impressions generallyshow little evidence of either the shell or of shell replacementmineralization as, for example, in the Geneseo and Middle-sex Shales. Exceptionally solid specimens occur either as cal-citic replacements and infills in calcareous nodules, as insome levels of the Rhinestreet Shale, or as pyritic internalmolds, as rarely in the Angola Shale. Associated benthicfauna is extremely limited, consisting largely of small lingu-loid brachiopods. The shales often appear rich in microflora.

Interpretation of this facies is part of the much-discussedproblem of the American Paleozoic black shale province asa whole. As suggested earlier (House, 1975b: 477), excessivedepths of formation seem unlikely. Sutton (1963: 97) gavedepth estimates of probably several hundred feet and possi-bly several thousand feet for the formation of the blackshales of New York. Lower figures, however, seem preferablefor the following reasons: (1) The generally cratonic settingof the facies, apart from the Michigan Basin, make depths ofseveral thousands of feet difficult to conceive on geophysicalgrounds. (2) The increasing evidence of unconformitieswithin the westerly black shale type regimes (Rickard, 1975;Baird & Brett, 1986a, b; Ettensohn, 1994) indicate eleva-tion and erosion more readily associated with shallownessthan extreme depth. (3) The evidence of winnowed and re-manié levels, such as the North Evans Limestone, is more

readily interpreted as part of a shallow setting. The settingsuggested is one in which the sediment/water interface isbelow the wave agitation zone and sufficiently below thethermocline to give euxinic and anaerobic conditions withinthe muds. Essentially these are infill deposits following deep-ening events.

References cited by Brett & Ver Straeten (1997) andMurphy et al. (2000) provide access to the vast literature onthe relations of tectonics, sea level. and water depth in thedeposition of black and gray shales in the AppalachianBasin. Particularly relevant to this study is the work of Mur-phy et al. (2000), a fine-scale biogeochemical analysis of coresamples through the facies of the upper Hamilton and lowerGenesee Groups in New York. They proposed a model inwhich the enhanced organic carbon burial in black shaleunits (like the Geneseo Shale) was the result of high surface-water productivity resulting from recycling, from the bot-tom to the surface, of the limiting nutrient phosphorus, afeed-back mechanism in a shallow marine ecosystem. Theirinterpretation does not support water-column stratification(i. e., anoxic conditions under a pycnocline) but rather limitsthe hypoxic conditions (a critical limiting factor for the ben-thic biota) to levels only at or near the sediment/water inter-face. The elevated productivity would enhance the ecologicalopportunities for groups like goniatites that occupied nicheshigher in the water column. Following their model, the rar-ity of goniatites in black shales would largely be the result oftaphonomic removal.

(2) Gray to Olive Shales and Mudrock.–As with theblack shale facies, there is a considerable range of color rep-resented in this wide group. Variations of gray are common-est. Olive-green coloration characterizes much of theCashaqua Shale facies of the classic Naples type, but alsotends to enter with increasing siltiness. There are transitionsalso into the black shale colors.

Goniatites are not rare in this facies, and are in placescommon. Generally they occur in a characteristic preserva-tion type with calcitic inner whorls and crushed outerwhorls. Of course, it is in these shales that the goniatite-rich

Text-fig. 18. Illustration of facies types and their inferred setting, largely based on the Upper Devonian Sonyea Group in New York (partlybased on Bowen et al., 1974).



concretions occur, but these represent a different seafloorenvironment and are treated separately. Nevertheless, the oc-currence of solid, calcitic inner whorls probably representsthe nascent development of the concretion facies. It is no-ticeable that crushed remains are generally absent.

In the facies transect of Text-fig. 18, bioturbation andevidence of trace fossils enter in with the gray shale facies ofthe basin and open shelf. Associated faunas, excluding thoseof the concretions, include commonly abundant occurrencesof bivalves such as Pterochaenia, rarer Buchiola, and othersmaller forms. These might well have been epiplanktonic(House, 1975b: 482; Grimm, 1998). At certain levels in theGenesee Group, the top surfaces of dark gray shales towardblack have exceedingly abundant accumulations ofLeiorhynchus and Barroisella (Thayer, 1974). Ager (1967)supported an epiplanktonic role for Leiorhynchus in this fa-cies; this interpretation has its origins in the classic essay ofSchmidt (1935). Thompson & Newton (1987), however,reinterpreted the Leiorhynchus clusters in the Geneseo Shaleas opportunistic epifaunal assemblages inhabiting restricteddysaerobic environments at the pycnocline/slope boundary.The relative rarity of shelled benthos is still noteworthy.

(3) Fine Siltstones.–Clastic tongues of well-bedded fine-grained siltstones, usually well cemented, range in colorfrom light gray to olive; dark colors are unusual. These unitstend to characterize the upper, coarser parts of majorrhythms or cycles and, more unusually, minor rhythms. Forthe major rhythms, they suggest progradation in the finalstages of basin infill when little accumulation space remains.Often substantial thicknesses are involved (Text-fig. 3). Frag-mental shell-bearing siltstones occur at some levels and theseseem to be facies intermediates approaching that of siltysandstones and sandstones discussed briefly below in whichrich shelly faunas often occur.

In this facies, goniatites are often common but tend tobe restricted to the bases of siltstones. Specimens include thelargest known of certain genera, notably of mantiococeratidsand triainoceratids, and, particularly in the Gardeau and En-field, respectively. Specimens are preserved solid or withsome crushing of outer whorls essentially as siltstone infillswithout calcitic inner whorls. Early diagenetic matrix ce-mentation is suggested.

The limited fauna of this facies is mostly associated withthe goniatites and takes the form of shell incrustations oftenindicating growth on dead shells on the sea floor. Large spec-imens of orbiculoid brachiopods (Clarke, 1899a; 1899b: pl.3, fig. 4) growing on the inside of large body chambers areknown; encountered more frequently are growths of aulo-poroid corals in the same position within empty body cham-bers. This suggests that this facies is approaching a betteraerated environment and represents the entry of calcitic-

shelled benthos. Burrowing in these siltstones is rare.(4) Turbidites.–Some, if not many, of the siltstones dis-

cussed above could be turbiditic in origin, but some are iso-lated turbidite beds within shale facies (see 1 and 2 above).Some turbidite beds have been traced for considerable dis-tances.

At the mouth of Farnham Creek, Lake Erie Shore (Loc.72, Text-fig. 15) is one such turbidite, approximately 20 cm(8 in) thick, within the seventh microcyclothem or rhythmfrom the base of the Angola Shale (West Falls Group). Thisturbidite unit, varying little in thickness, has been traced atthe same level for 50 mi (32 km) to the east (occurrencesare shown on Text-fig. 15). This can be referred to as the“Farnham Creek Bed.” No goniatites are associated with it.

A similar occurrence is the Bluff Point Siltstone, a 1.5-11 cm (0.6-4.3 in)-thick convolute-bedded unit seen withinthe gray shales of the West River Shale (Genesee Group) atmost localities between the Genesee Valley and Seneca Lake.It is a level that can be traced farther to the east and west(deWitt & Colton, 1978: pls 2-6). The goniatites (Koenen-ites and Lobotornoceras) occur in the cycle below it.

The Crosby Sandstone is a marker bed at the contact be-tween the Penn Yan Shale and overlying Ithaca Shale andSandstone in the vinicity of Keuka Lake; there calcareousconcretions in its base contain the goniatites Koenenites andTornoceras. As noted in the stratigraphic section above, theCrosby correlates west to Canandaigua Lake to the “hiatusconcretion horizon” of Baird (1976) in the Upper Penn Yanand eastward as a prominent siltstone within the Ithaca tosections in the Seneca and Cayuga valleys (de Witt &Colton, 1978: pl. 3). The Crosby is a complex, bioturbated,multi-event bed, the probable turbidite origins of which areobscure.

In the Namurian of Devon, England, goniatite nucleiare commonly found with broken shells of large specimensat the base of similar turbidite units (Pinkhoe, Brickpit, Ex-eter). The New York occurrences differ in that the goniatitesare large and commonly complete. It could be that they rep-resent shells swept down from higher levels on the paleos-lope with the onset of a turbidite flow.

(5) Silty Sandstones and Sandstones.–In rocks of this fa-cies, which present more clastic rocks to the east, goniatitesare virtually unknown apart from isolated occurrences. Thisis the facies that embraces the delta platform, delta front,and pro-delta facies described, for example, by Bowen et al.(1974). A very rich shelly fauna occurs here that is well doc-umented as the classic Naples Fauna but does not concernthis discussion. Yet farther landward, alluvial facies yield nogoniatites at all despite the evidence that the shells of theliving Nautilus are often blown or swept well onshore(House, 1987).

Carbonate Facies.–TheTully and Genundewa represent theonly significant limestone tongues in the part of the se-quence dealt with here. The latter, because of its distinct na-ture warrants individual comment. Otherwise, carbonateoccurrences are localized, taking the form of concretions ofKnollenkalk and Kramenzelkalk facies. The general absenceof a carbonate facies belt in the later Devonian of New Yorkis largely responsible for giving it such a distinctive overallcharacter, in sedimentological, paleontological, and paleoe-cological terms.

(1) Limestones.–Only the Tully Limestone truly belongshere and this has been the subject of detailed treatment inHeckel’s (1973) magnificent monograph and more recentlyin Baird & Brett (2003) and Baird, Brett & Bartholomew(2003). As has been documented earlier, only one levelwithin the Moravia Bed or West Brook Member (Cooper&Williams, 1935) has yielded goniatites.

The Tully Limestone level has been regarded as trans-gressive since the late nineteenth century (Frech, 1897: 255),a view confirmed by the work of Cooper (1968), Cooper &Williams (1935), Cooper et al. (1942), and Heckel (1973,1997). Formerly it was regarded as the Upper Devoniantransgression, but the name Taghanic Onlap (Johnson,1970, 1989) is appropriate because the actions of the Sub-commission on Devonian Stratigraphy (SDS) and Interna-tional Union of Geological Sciences (IUGS) have relegatedthe level to the Middle Devonian (Klapper et al., 1987). Itmight be expected that the limestone phase of majorrhythms would represent the shallowest facies, but it is bestexplained by deepening, for the following reasons. (1) In thegeneral paleogeographical setting of the Devonian Ap-palachian Gulf, clastic facies predominate and the TullyLimestones will represent a period when clastic depositionwas far distant; this is most easily achieved by sea-level rise.(2) The entry of “exotic” elements, such as harpid trilobites,the brachiopodHypothyridina, and the goniatite Pharciceras,indicate links to world oceans and ready migration, such asis best explicable by deepening. (It should be noted thatHy-pothyridina venustula (Hall, 1867) was assigned to a newgenus, Tullypothyridina, by Sartenaer in 2003.) (3) In theNew York facies transect, as demonstrated by Cooper &Williams (1935: text-fig. 3), Johnson & Friedman (1969),Rickard (1975, 1981), Baird & Brett (2003), and Baird,Brett & Bartholomew (2003), the Tully facies tongues farto the east, carrying marine neritic facies within successionsthat also bear the Gilboa Forest in a way most simply ex-plained by transgression. That is not to suggest that greatwater depths are required to explain the Tully Limestonefaunas, but transgression clearly led to the facies change.Westward, the thinning of rock units, and the evidence ofunconformities (Text-fig. 7) both below, within, and after

the Tully, shows that intermittent seafloor elevation led towinnowing and erosion in those areas as they had in theHamilton. Baird & Brett (2001, 2003) and Baird, Brett &Bartholomew (2003) have differentiated parts of theTully ashighstand and lowstand deposits. They recognize a majorlowstand event in the Upper Tully marked by a regional dis-conformity at the base of the Bellona Bed in western NewYork and the base of the West Brook Shale in eastern NewYork. This lowstand marks the reestablishment of theHamilton Fauna (brachiopod-coral biofacies); the subse-quent deepening (West Brook Bed-Moravia Bed) is markedby the incursion of Pharciceras and the highstand deposits ofthe Fillmore Glen Bed and post-Tully Geneseo Shale.

After the Tully Limestone, good limestones are not de-veloped higher in the Devonian of New York, but a similarinterpretation applies to the limited limestones of the Lodiand Genundewa. Both of these are characterized by theabundance of ammonoids (respectively, Ponticeras and Koe-nenites especially) again indicating a link with the worldoceans. The styliolinid facies of the Genundewa is a pelagicdeposit, but the styliolines, thought to be planktonic, couldwell have been concentrated by drifting. No great depthneed be required and the vertically orientated goniatites inthe Lodi (Pl. 4, Fig. 5) suggest relatively shallow water. Theinterpretation of theTully, Lodi, and Genundewa limestonesas relatively deep in relation to the clastic facies has been il-lustrated in recent sea-level curves for New York (House &Kirchgasser, 1993; House, Menner et al., 2000).

(2) Large Concretions.–Characteristically developed atspecific levels within the shale facies are concretionary hori-zons. The colors of the concretions tend to match the colorsof the shales in which they occur. The larger concretions,taken arbitrarily as those over 20 cm in diameter, are consid-ered here. Usually these are depressed-ovoid in vertical sec-tion and circular in plan. Some larger examples showinterpenetration of sphaeroids. Concretions are usually sep-arated by distances in excess of their width along specifichorizons. Rarely they coalesce as concretionary limestones,of which the Corell’s Point Goniatite Bed (Loc. 97) is thebest example, which can be traced inland from the shore ofLake Erie. Some of the more spherical concretions in theGenesee Group reach over a meter in diameter, and moredepressed examples in the Rhinestreet Shale near Lake Eriereach more than twice that size.

The largest concretionary levels in minor rhythms or cy-cles (Text-fig. 19C) tend to occur in the upper part of therhythm or cycle, below the black shale that introduces thesucceeding unit. In general, there is also a similar relation inthe major rhythms or cycles. For example, larger type con-cretions occur in the upper part of theWest River Shale andlower Cashaqua Shale on either side of the intervening Mid-


dlesex black shale. Other large concretions occur within theuppermost part of the Cashaqua Shale and upperRhinestreet Shale, on either side of the main Rhinestreetblack shale tongue. The largest concretions of the HanoverShale occur a little above the black shale tongue of the PipeCreek Shale. As will be discussed later, for the majorrhythms, this introduces an element of cyclicity into the pat-tern.

Concretions tend more frequently to be septarian wherethe surrounding shale is darker gray or blacker in color. Atcertain levels, and perhaps areas, the normal crystalline cal-citic infill has been replaced by barite, and fossil shells havealso been so replaced. The exceptionally fine development ofthis type is seen in the Shurtleff Septarian Horizon (Text-fig. 10); other levels will be referred to in the section onpreservation. The goniatite beds we refer to as “Fossil LogHorizons” (FL) are concretionized rafts of driftwood inwhich the plants and shells accumulated as hydrodynami-cally similar, lightweight materials as suggested by Maeda &Seilacher (1997).

Fine, solid, goniatite specimens, often difficult to extract,are common in many of the large concretionary horizons, al-though perhaps rarest in the largest. Commonly there is anassociated fauna of gastropods and bivalves, but these areeven more difficult to extract, being smaller and mostly re-crystallized replacements. At baritized levels, where acid dis-solution enables easy preparation, a great variety of formsare known, as described by Clarke (1899a, b) from theShurtleff Septarian Horizon (Loc. 41). It is interesting thatwhere concretions start to coalesce, forming essentially aconcretionary limestone bed, as the Genundewa Limestoneand Corell’s Point Goniatite Bed, gastropod and bivalve fau-nas increase both in size and abundance. The GenundewaLimestone is distinctive in other respects, however, and isdiscussed separately below. In the Corell’s Point GoniatiteBed, the numerous gastropods are often similar in size to,and roughly homeomorphic with, the associated Cheiloceras.

Because shells in these concretions are uncrushed, it isapparent they represent early diagenetic cementation eventsperhaps arising from nucleation centers associated with or-ganic remains. The availability of calcium carbonate for theprocess, however, indicates that the concretionary levels rep-resent different environmental conditions than that of thebackground shale above and below. The baritization isthought to be a much later epigenetic event.

(3) Small Concretions.–There is a series in size of con-cretions from those just considered down to small concre-tions that are essentially nodules approximately 10 mm indiameter. Levels with very small concretions are rare. Thelarger among this group are similar in other characters tothose already described, and occur in a similar position

within minor rhythms (Text-fig. 19C); these can rarely beseptarian. Their color is similar to the shales in which theyoccur. Generally, with decreasing diameter, there is a trendto greater sphericity.

The smallest concretions often show a pyritic nucleusand evidence that a burrow or other organic center occurswithin them. In places, as not uncommonly in the CashaquaShale, the nodule can comprise a goniatite body chamber,but usually not the phragmocone. In smaller concretions,evidence of nucleation of the concretion around central or-ganic material is more often seen than in larger concretions,even those containing goniatites.

This group of concretions appears to have formed inprogressively deeper and less calcareous sedimentary envi-ronments than the large concretions, but this is an overallimpression and not universally tenable. Where goniatites arepreserved, they are usually solid and indicate lithificationbefore any appreciable compaction had taken place.

As with the larger concretions, molds of goniatites withrecrystallized shells are sometimes replaced by barite or,more rarely, silica. Examples of this type of small noduleswith diagenetic barite rosettes occur in the lower HanoverShale (Pepper et al., 1985). This replacement will have takenplace long after concretion formation.

(4) Knollenkalk and Kramenzelkalk Levels.–Deeper waterDevonian deposits of the European Hercynian Facies com-monly comprise micritic nodular limestones and cephalo-pod limestones known as Cephalopodenkalk, Knollenkalk, orgriotte in the European literature. This represents a facies inwhich goniatites occur abundantly at appropriate levels. Inplaces, the more blotchily colored nodular limestones havea distinct “ant-eaten” appearance in weathered outcrops.This gave rise to the term Kramenzelkalk for a distinctive fa-cies in which the carbonate nodules and enveloping shalesare differentially weathered with the nodules hollowed toform a reticulate network of cells. Tucker (1973) consideredthese facies types as pelagic limestones, a term more geneti-cally precise than the facts warrant.

In New York, at certain levels in the Frasnian, nodularlimestones of a similar sort occur. These are commonly redand green in coloration, form distinct if not thick beds, andshow a curious weathering. Generally they are coarser ingrain size and more impure than typical European types,even if their characteristics bring them within the termino-logical range of European usage.

This facies seems to belong to a shallower environmentthan those so far discussed, but which is unusual in being as-sociated with rises on the sea floor. The European examplescharacterize the crests of substantial seafloor elevations orSchwellen. The New York examples suggest a similar, if lessspectacular, setting.



The Parrish Limestone, within the Cashaqua Shale(Text-fig. 10, Beds 47/2 and 48/1), occurs on the crest of aclastic wedge termed the Rock Stream Siltstone, and appearsto owe its characteristics to the oddly raised environmentpresented by the crest of the wedge. The Parrish Limestone,described by Clarke (1885: 39) as a Kramenzelkalk facies, isa famous source of manticoceratids in the region close toNaples (Clarke, 1885; Kirchgasser 1965, 1975). Passing lat-erally from this region, both to the west and to the east, thefacies changes to shales with nodules such as those previouslydiscussed.

Two other examples occur higher in the sequence (Text-fig. 3): levels in Angola Shale equivalents near Varysburg(Wyoming County; Text-fig. 15, Loc. 79/10), and on John-son Creek (Wyoming County; Loc. 81/10), which are thindevelopments of this type of facies. At both localities thereare other levels rather higher but well below the horizons of“TheTrinity” black shales. A notable one with goniatites oc-curs within the West Hill/Nunda interval at Varysburg(Gassman Road) (Loc 80/14). Other levels occur in thelower part of the Hanover Shale, as inWalnut Creek (Chau-tauqua County; Text-fig. 16, Loc. 89a), and red and greenKramenzel-type nodular horizons are found in the upperHanover Shale in Beaver Meadow Creek, Java (Loc. 92).Also, as it passes eastward, the Corell’s Point Goniatite Bedappears to lose its concretionary nature, and specimens oftypical Cheiloceras from the Java region (Wyoming County)appear to have come from nodular limestones; it could bethat the facies recurs higher in equivalents of the GowandaShale.

(5) Styliolinid Limestones.–Quite an unusual and sepa-rate facies is represented by calcareous levels where the smallconical shell of Styliolina fissurella (Hall, 1845), and perhapsother dacryoconarids, become so important as to form dis-tinct units. The best developed level of this sort is the Ge-nundewa Limestone, but other levels of a similar sort occurin the Penn Yan Shale below and theWest River Shale above.Recognition of the Genundewa Limestone by styliolinid oc-currence alone is most untrustworthy. A black shale beneathit (Text-fig. 8) is used here as the most accurate means ofdefining its base.

Whereas the preceding carbonate facies seem to be re-lated to particular depth and site-related factors, the Styli-olina facies relate to a peculiar abundance at particular timesof an organism that is thought to be planktonic and usuallypelagic and whose remains fell to the sea floor after death(Fisher, 1962; Lindemann, 2002). Thus their accumulationprobably depended on factors of abundance, wind direction,tidal patterns, current movements, and other factors ratherthan on seafloor characteristics. Commonly associated withthe styliolinids in the Genundewa are small snails, and it has

been suggested (House, 1975b) that these might have beenplanktonic bubble-snails, associates in the ecology of the liv-ing Styliola. Goniatites occur abundantly at certain levels inthe Genundewa, as at Bethany Center (Loc. 24b), togetherwith bactritids and orthocones in a manner that suggestsdrift accumulation. But there is also a substantial benthonicfauna (Clarke, 1904; Sass, 1951) that probably flourishedin the accumulating styliolinid sands.

Styliolinid limestones with goniatites are developed tothicknesses reaching 50 mm at levels in the late Penn YanShale, and West River Shale and correlative horizons, in-cluding the Genundewa, have been found in the HarrellShale in the region near Landes Post Office, West Virginia(House, 1978), and in the Harrell Shale in central Pennsyl-vania (Kirchgasser, 1996b). The Squaw Bay Limestone inMichigan is a similar facies. Such widespread styliolinidunits with goniatites are interpreted as trangressive phases.Styliolinid limestones or concretions, often with goniatites,occur at several levels in the Sonyea Group and West FallsGroup. Dacryoconarids (sensu Fisher, 1962, and Linde-mann, 2002) range into the upper Hanover Shale, and ho-moctenids have been found at the top of the Hanover Shaleand in the lower Dunkirk Shale (Yochelson & Kirchgasser,1986; Over, 1997a, b; Over et al., 1997).

MODE OF GONIATITE PRESERVATION

Goniatites in the New York Devonian can be preserved in awide range of ways but a few types are typical. These arelargely a result of the variety of facies types in which theyoccur and hence reflect the classification of facies already re-viewed.

No goniatites preserving the original aragonite shell ma-terial are known in the New York Devonian. The shells, ifpreserved, are invariably recrystallized, normally as calcitebut also, rarely, as barite, as with the material of the ShurtleffSeptarian Horizon (Loc. 41/1) in the upper Cashaqua Shalein Livingston County. Larger specimens usually lose eventhis, and such shells are represented by external and internalmolds. Attention should be drawn to the fact that the exqui-site material of the inner whorls of goniatites, especially thatfigured by Clarke (1899a, b) and refigured here (for exampleon Pl. 21) was exposed by Clarke by dissolving away recrys-tallized shells set in mudrock and, when dry, making guttapercha molds. The resultant preservation is incredibly finebut the original specimens were not retrieved.

The commonest of the natural internal molds are thosepreserved in pyrite. These rarely exceed 10 mm in diameterand are associated with dark gray or black shales, as in theWest River Shale and Rhinestreet Shale. Remanié accumu-lations of pyritic material, as in the Leicester Pyrite (Text-fig.7) and other levels are concentrates resulting from the win-

nowing and erosion of shales which formerly contained thepyritic molds (Baird & Brett, 1986a).

CAUSES OF MAJOR AND MINOR RHYTHMS AND CYCLES

As has been illustrated (Text-figs 3, 8, 13, etc.), transgressivepulses, especially those initiated by black shales, either at amajor or minor rhythmic scale, form a characteristic featureof the stratigraphy of the late Devonian in New York State.Most are clearly of a rhythmic type, with an asymmetricalpattern ABCABC or abcabc (Text-figs 19A, C). As notedabove, Murphy et al. (2000) have proposed a within-basinrecycling model in interpreting the geochemistry of blackand gray shale cycles in the upper Hamilton and lowerGenesee Groups. As has been deduced from previous con-sideration, the ABC, abc sequences are interpreted as shal-lowing upward sequences with the shallowest facies (C)abruptly followed by deepest facies (A). In contrast, themajor cycle of the Sonyea Group with a pattern ABCBA(Text-fig. 19A) is a true symmetrical cycle. In this case, theshallowest facies (C) is succeeded (gradually) by deeper waterfacies (B) which in turn is followed by the deepest facies (A).

Possible causes of such rhythmicity of cyclicity have beendebated since they were recognized as features of the Jurassicrhythms in the South of England by Conybeare & Phillips(1822). For the smaller scale rhythms, usually of the orderof 0.5-2.5 m, a cause from climatic changes would dependon changes in outer atmosphere insolation as controlled bythe changing orbits due to perihelion, precession, and theshort and longer eccentricity cycles. The resultant climaticshifts could cause small sea-level changes due to simple ex-pansion, the freezing and melting of polar and mountainice; on land such changes of climate would modify vegeta-tion cover, rates of weathering and erosion, and the natureof sediment reaching the basin. The New York sequencewith which we are concerned shows no long-continuedsmall-scale rhythmicity as does, for example, the Devonianpelagic facies of the Montagne Noire (House, 1995). Small-scale (meter-scale) rhymicity occurs in parts of the New Yorksuccession, most notably in the West River Shale (thetwenty-three rhythms) and in the lower Angola and lowerHanover shales as discussed in the stratigraphy section. Nev-ertheless, there are too many transgressive pulses perhaps

Text-fig. 19. Patterns of sedimentary rhythms in late Devonian rocks of New York State. All scales approximate (from House & Kirchgasser,1993). A. Diagram simplifying data on facies within the Sonyea Group of western New York indicating the sequence in one type of majorrhythm or cyclothem; Middlesex Shale at base. B. Diagram simplifying data on facies within the lower West Falls Group of western NewYork indicating the sequence in another type of major rhythm or cyclothem; Rhinestreet Shale at base. C. Diagram indicating the sequencein a typical minor rhythm or microcyclothem based on examples within the Angola Shale of the West Falls Group.


due to other causes, such as tectonics, which interrupt sed-imentation, to make the New York succession easy for se-quence or cyclostratigraphic analysis. Primary evidence forthe operation of Milankovitch cycles is therefore unlikely tobe forthcoming, and interpretation must be by analogy withthose parts of the stratigraphical column now well docu-mented in Milankovitch and oribital forcing terms.

In the New York setting, interpretations of larger scalerhythms fall into two main groups. First are those invokingintermittent basinal subsidence probably caused by periodicmovement on faults bounding depositional basins associatedwith convergent-margin tectonism (the tectophases of Et-tensohn, 1985, 1994). Second are those invoking global eu-static rises of sea level (see references below). Theories of thelatter group would accept the importance of differentialbasinal subsidence in explaining the pattern of thicknessvariations in the New York Devonian, but would considerthe subsidence to be generally gradual and not intermittent.

The discrimination between these groups of hypothesesreally depends on whether transgressive pulses can be corre-lated over wide areas of the globe in Devonian time. It isthought that for the group of strata with which we are con-cerned there is increasing evidence that this can be sustained,and that the second group of hypotheses seems generally tohave been the more important. Increasing evidence in favorof the latter group of hypotheses includes the observationthat many of the major facies movements are internationalin scope (House, 1983, 1985; Johnson et al., 1985, 1986),but especially the detailed correlations of the Frasnian inWestern Australia (Becker et al., 1993), northern EuropeanRussia (House, Menner et al., 2000), and New York (House& Kirchgasser, 1993)

The main transgression indicated by theTully Limestoneand especially the West Brook Member or Upper Tully di-visions (Heckel, 1973; Baird & Brett, 2001, 2003; Baird,Brett & Bartholomew, 2003) has been widely recognized inNorth America as the Taghanic Onlap by Johnson (1970),which is just a new name for the American Upper Devoniantransgression of the consistent literature of earlier scientistsdating back a century. Internationally, the importance ofonlap movements at this time-interval have been docu-mented by House (1975a). It is now recognized, however,that in New York, several phases of transgression are in-volved (Text-fig. 7) and much has still to be learned approx-imating these, both with regard to sea-level changes andextinctions, internationally as well as locally in New York.Depophase IIa of Johnson et al. (1985) commences at thebase of the Tully Formation.

The Genundewa has also been interpreted as a transgres-sive level in New York (House, 1983, 1985; House & Kirch-gasser, 1993; House, Menner et al., 2000). The level

approximately corresponds to the lower Assise de Frasnesonlap of Europe. Again the contrast between the lower andupper parts and the preceding black shale indicate that thisis also a polyphased event. It corresponds to depophase IIbof Johnson et al. (1985).

The Middlesex black shale pulse seems to correspondwith the deepening that resulted in the growth of BelgianF2d reefs (House, 1985). The same level appears to be rec-ognizable as the base of the Domanik in northern EuropeanRussia (House, Menner et al., 2000) and the same pulse ofdeepening has been recognized inWestern Australia (Beckeret al., 1993). This is the base of depophase IIc in the termi-nology of Johnson et al. (1985).

The major deepening event represented by theRhinestreet black shale can similarly be correlated in otherareas, for example with the initiation of the F2h reefs in theArdennes (House, 1985). This appears to be the base of de-pophase IId in the terminology of Johnson et al. (1985)whose major depophase boundaries were linked to the groupterminology of Rickard (1964, 1975), in this case the baseof the West Falls Group. But some (Sandberg et al., 2002)seem to be taking the base at the later semichatovae trans-gression, which could correspond to one of the later trans-gressive pulses in the Rhinestreet recognized by Sutton(1960, 1963), but this interval is still poorly documentedby conodonts in New York (Klapper et al., 1995).

The Pipe Creek black shale is a widely recognized inter-national transgressive event (dysoxic pulse) that correspondsapproximately to the German Lower Kellwasserkalk trans-gression and the initiation of the deep-water Schistes deMatagne in Belgium (Schindler, 1990, 1993; House, 1985,2002). No zonable conodonts are known from the PipeCreek but the level is probably within Zone MN12 (?Upperrhenana) (Over, 1997a, b; Klapper & Becker, 1999). TheKellwasserkalk transgressions in Europe were first discussedin detail by Buggisch (1972) and recognition was then ex-tended to North Africa by Buggisch & Clausen (1972). Thecorrelation of the Pipe Creek Shale with the Lower Kell-wasserkalk of Germany was not recognized by Johnson et al.(1985, 1986), who correlated the base of the RhinestreetShale with the Lower Kellwasserkalk, a view now known tobe in error.

Through the work of Over (1997a, b, 2002; Over et al.,1997), much more detailed correlation is now available forthe placing of the newly defined Frasnian/Famennianboundary (Klapper et al., 1987). It clearly precedes the baseof the major tongue of the Dunkirk Shale (boundarymarked in Text-fig. 16). Thus the underlying black shale,known informally by John Huddle as the Rider Bed (fromthe miner’s term for any underlying or overlying leaf from acoal seam), seems to be the New York equivalent of at least


part of the Upper Kellwasserkalk. In view of the detailed doc-umentation of the phases of the Kellwasser Events(Schindler, 1990, 1993), work is still needed to give precisedocumentation with Europe and other areas (House, 2002).

The Dunkirk black shale pulse corresponds to the Ne-hden transgression of Germany and other areas that initiateCheiloceras Stufe faunas, which was named the NehdenEvent by House (1985), but Cheiloceras arrives later in NewYork. Although the Dunkirk Shale corresponds to the Ne-hden Event (Becker, 1993a) and transgression of the earlyFamennian of Europe, relevant goniatite faunas do not ap-pear until the Gowanda Shale. The Dunkirk corresponds tothe base of depophase IIe in the terminology of Johnson etal. (1985, 1986).

These main events are interpreted as transgressions andmost are associated with a flooding of dysoxic- to anoxic-type sediments. It is not to be expected that the sedimento-logical effects of transgressions will be exactlycontemporaneous in different areas and where those areascan be related to different positions on the hypsometriccurve.

The increasing transgressions through the Frasnian havebeen argued as the main cause of late Frasnian extinctions,and indeed, these are mostly accomplished by the Kell-wasserkalk transgressions (House, 1975b). It is not thoughtthat offlap and regression leading to “perched faunas” (John-son, 1974) are a reasonable interpretation because the mainrecognizable regressive events, in Europe as in North Amer-ica, do not appear until the late Cheiloceras Stufe, well afterthe extinctions.

Although major global eustatic changes of sea level canexplain similar major rhythmic patterns in different conti-nents, a similar interpretation for microcycles cannot betested because correlative tools are too imprecise. The loworder of thickness involved in the minor patterns (Text-fig.19C) could be equally well explained in sedimentologicalways alone by migrating distributaries and other mecha-nisms against a general background of basinal and basin-marginal subsidence.

If the global eustatic cause for the major rhythms comesto be accepted, then the question of a mechanism for sea-level changes is required. Two of the main contenders arechanges in the amount of polar ice and volumetric changesat constructional plate margins. The glacio-eustatic inter-pretation is difficult to sustain in the absence of clear evi-dence for significant polar ice in the Devonian and thegenerally warm and equable climate generally inferred in aperiod that represents the maximum in faunal and floral di-versity in the Paleozoic. Other contenders for the sea-levelchanges include orbitally-forced climate changes (House,1985; 2002) and comet-shower impacts (Sandberg et al.,

2002).

EVOLUTION OF LATE DEVONIAN GONIATITES

A current view of the origin and radiation of Devonian go-niatites and their Late Devonian derivatives, the clymeniids,is illustrated in Text-fig. 20. This also shows the “events,”usually marked by widespread international transgressionand anoxia, which apparently had a major role in controllingevolution of the group (House, 1985, 1996; Becker, 1993b;Becker & House, 2000). The group was derived from or-thoconic Bactritida and the coiled goniatites with a dorsal si-phuncle first appear a little above the base of the Emsian.By the beginning of the late Givetian (formerly basal UpperDevonian), at the Taghanic Event, most earlier Devonianstocks had become extinct, including the families Agoniati-tidae, Pinacitidae, Sobolewiidae, and almost all the Maenio-ceratidae.

Formerly, all the Frasnian gonitiates apart from the An-arcestidae and Tornoceratidae were grouped together as theGephuroceratoidea. It is now known, however, that thePharciceratidae and its derivatives (Eobeloceratidae, Pettero-ceratidae, Triainoceratidae, and the problematic Devono-pronoritidae) have their origin in the Maenioceratidae bythe proliferation of umbilical lobes (Bensaïd, 1974).

Unless otherwise noted, the framework of classificationadopted here corresponds to that adopted generally in recentyears (House, 1981a; Becker & House, 1993, 1994a, b;Becker, 1993b).

Ranges of genera given here are based on internationalwork resulting from investigations since 1954. More recentare the detailed investigations resulting from work with theSubcommission on Devonian Stratigraphy in the definitionof stage and series boundaries and in relation to those“events” that appear to have significantly affected the evolu-tion of the group. The most significant ones are marked inText-fig. 20. This work has been greatly aided by the preciseconodont biostratigraphy established for the late Devonian(Ziegler, 1962, 1971; Ziegler & Sandberg, 1990; Klapper,1989, 1997, 2000; Klapper & Becker, 1999) that has givenevidence of age independent from the goniatites themselves.Text-fig. 21 is a range diagram for late Devonian taxa knownworldwide over the period with which we are concerned inNew York – the late Givetian to the mid-Famenian. Manyof these genera are not known in New York or in NorthAmerica.

BIOSTRATIGRAPHY

INTERNATIONAL GONIATITE ZONATION

OF THE LATE DEVONIAN

Detailed work on the ammonoid zonation of the Devonian


Text-fig. 20. Diagram illustrating the evolution of Devonian ammonoid families. The width of bars corresponds to the number of generaat particular times as indicated by the scale. The timescale used is based on Becker & House (1994a) and Becker (1993b). Also indicatedare the named environmental stress events. Modified from House (1996).


Text-fig. 21. Diagram illustrating the ranges of late Devonian goniatite genera internationally. From Becker & House (1994b); Famenniandata from Becker (1993b).



essentially began in Germany when Devonian studies fol-lowed the celebrated work of Albert Oppel (1831-1865) onthe Jurassic. Fritz Frech (1861-1917), August Denckmann(1860-1925), Emmanuel Kayser (1845-1927), RudolfWedekind (1881-1961), and Hans Matern (1903-1933)were chief among early workers. In more recent times no-table contributions on the Famennian were made by H.Schmidt &O. H. Schindewolf, B. I. Bogoslovsky, D.Weyer,J. D. Price, D. Korn, and R. T. Becker.

As noted by Becker & House (2000: 115), the potentialof ammonoids for international correlation within the De-vonian is limited by the demonstration in recent decadesthat synchronous but widely separated sedimentary basinscan contain different sequences despite the pelagic lifestyleof the group. The faunal distinctions appear to be the resultof many factors including the individual structural and facieshistories of basins, the limitations of faunal exchange be-tween open-marine environments by paleogeographical bar-riers resulting in endemic taxa, sedimentary discontinuities,and discontinuous representation of ammonoid biofacies.Nevertheless, using the wealth of new data generated by am-monoid workers from many sections around the world,Becker & House (2000) proposed a new and much refinedzonation that is correlated with the celebrated conodontzonation and with the established series and stage bound-aries.

Because the major early contributions on late Devonianammonoid biostratigraphy were based on sequences in Ger-many, it is appropriate to review that which has been termedthe “standard” succession established by the work of Frech,Denckmann, Wedekind, Schindewolf, Matern, Schmidt,Lang, and others in the Rheinish Schiefergebirge (RhenishSlate Mountains). A brief account of the characteristics andgeographical distribution of Devonian ammonoid zones andfaunas has been published (House, 1979) and need not berepeated here. It is important to bring this up to date bynoting especially the elevation of the lunulicosta Zone (for-merly included in the Manticoceras Stufe) to a new Pharci-ceras Stufe (House, 1985) in the position below the nowmuch-raised conodont-defined Middle/Upper Devonianboundary (Klapper et al., 1987). The discussion that followsis a brief introduction to the data, methodology, and termi-nology of the international ammonoid zonation of Becker &House (2000).

The major subdivisions of Devonian ammonoid zona-tion are:

Upper Devonian (UD)Famennian

UD VI Wocklumeria StufeUD V Clymenia StufeUD IV Platyclymenia Stufe

UD III Prolobites StufeUD II Cheiloceras Stufe

FrasnianUD I Manticoceras Stufe

Middle Devonian (MD)Givetian

MD III Pharciceras StufeMD II Maenioceras Stufe

EifelianMD I part Anarcestes Stufe and Pinacites

StufeLower Devonian (LD)

EmsianLD III Anetoceras Stufe and part Anarcestes

StufePragian (LD II) No ammonoids known.Lochkovian (LD I) No ammonoids known.

For the Upper Devonian, the former Stufen or major di-visions or zones of Wedekind & Schindewolf are essentiallyretained, each designated by Roman numerals (UD I-VI)and the name of a genus. Although never explictly defined,the Stufen are essentially taxon range zones that are based onthe first and last occurrence of a single taxon. Wedekind &Schindewolf subdivided the Stufen using Greek letters; thusUD II, the Cheiloceras Stufe of the Famennian Stage, wasdivided into IIa and IIb. Recent work has added consider-able precision to the zonal subdivision of the Stufen. For theLower and Middle Devonian, the revised zonation usesRoman numerals and the Roman alphabet to subdivide theStufen (e. g., MD III-A through E for the Pharciceras Stufeof the upper Givetian Stage) (Becker & House, 1994b).

For the Frasnian Stage of the Upper Devonian, the re-vised zonation, based largely on ammonoid successions inthe Canning Basin, Western Australia, and New York, fol-lows the same methodology. The Manticoceras Stufe, UD I,is divided into twelve lettered divisions, UD I-A through L,which replace the four divisions (I alpha to I delta) ofWedekind. The new subdivisions for the Upper Devonian asa whole are shown in Text-figs 20-21 and for the Frasnianand Famennian in Text-fig. 22.

The divisions in the zonation of Becker &House (2000)are essentially interval zones, defined primarily by the entryof genera rather than the occurrence of specifically namedzonal species. The entry of genera is more appropriate forinternational use because each area that has been examined(New York included) has a regional zonation, the character-istic feature of which is the dominance of local (often en-demic) species or subspecies. The new internationaldivisions are thus defined as Ammonoid Genozones. Becker& House (2000) also introduced an idealized parallel

Text-fig. 22. Zonation of the late Devonian using ammonoids plotted against the conodont zones.Based on Becker & House (2000).


scheme of Ammonoid Standard Zones, based on the entryof species (zone fossils) with international or interbasinaldistribution but that might not occur in all basins with con-temporaneous ammonoid faunas. For regional successionswith less cosmopolitan or endemic forms, Becker & House(2000: 115) recognized Ammonoid Regional Zones, whichare useful for intrabasinal correlation but whose markerforms (genera or species) “…may differ from standard or in-ternational zones (for example endemics may be used) ormay have shorter ranges than known from the total globalrecord (regional topozones); for example, delayed appear-ance of Manticoceras in various areas in comparison to itsearly appearance in New York.” The broad key to the newzonation (Text-figs 20-21) shows the international range ofgoniatite families and genera in the Frasnian as currently un-derstood. The zonation of the succeeding Famennian is notindicated because clymeniids, which form the basis of thesubdivision, have yet to be found in New York State.

GivetianThe faunas of the Eifelian and earliest Givetian of New Yorkare not of concern here. A review of known records wasgiven by House (1962, 1981b) and Becker & House(1994b), but there is important new goniatite informationin the unpublished thesis of G. Kloc (1983) and in his pri-vate collections.

Maenioceras Stufe, MD II (A-D).–As has already been de-scribed in the section on the evolution of late Devonian go-niatites, the major break in the ammonoid sequence is attheMaenioceras Stufe/Pharciceras Stufe boundary, which wasformerly taken in Germany as the Middle/Upper Devonianboundary, and by most as the Givetian/Frasnian boundary(House, Feist & Korn, 2000). Toward the end of the Mae-nioceras Stufe interval, a wide range of typical Middle De-vonian goniatites become extinct, namely Sobolewia,Pseudofoordites, Wedekindella, Foordites, Holzapfeloceras,Cabrieroceras, Werneroceras, Agoniatites, and Sellagoniatites.All of these genera, except perhaps Foordites, Pseudofoordites,and Sellagoniatites, the precise ranges of which are uncertain,are documented in lateM. terebratum Zone faunas (MD II-C). For example, most of these genera occur in the Penton-warra Goniatite Bed of North Cornwall (House, 1963). Bythe boundary, therefore, the families Pinacitidae, Holzapfe-loceratidae, and Agoniatitidae become extinct, and the Mae-nioceratidae survive for only a very short time. This MDII-D extinction is the Taghanic Event of House (1985).Only a remnant of the Anarcestidae survives, perhaps to theearly Famennian, and gives rise to the Gephuroceratoideaand Pharciceratoidea. The Tornoceratoidea survived theTaghanic Event and appear to have given rise to all Famenn-

ian groups of the Ammonoidea (Text-fig. 20).

Pharciceras Stufe, MD III (A–E).–The succeeding lunuli-costa Zone (I alpha) of Wedekind (now Pharciceras Stufe,MD III) was first defined by Frech (1887: 433) using faunasfrom several localities in the Rhenish Schiefergebirge andHarz Mountains of Germany. The Pharciceras Stufe intro-duces members of the old suborder Gephuroceratina, first ofthe Pharciceratoidea and of the Gephuroceratoidea shortlythereafter. Because these groups are thought to be independ-ently derived from the Anarcestidae, a single suborder forthem is inappropriate.

The new subdivision of the Pharciceras Stufe recognizesfive divisions (Becker &House, 1994b, 2000). The fauna ofMD III-A is defined by the entry of Pharciceras and Mzer-rebites but with the hangover from the Maenioceras Stufe ofMaenioceras, Afromaenioceras, the anarcestid Atlantoceras,and tornoceratids, especially Tornoceras and perhaps Epi-tornoceras. MD III-B is defined by the entry of multilobedpharciceratids, especially Stenopharciceras [e. g., S. lunulicosta(G. & F. Sandberger, 1850)]. MD III-C shows the entry ofSynpharciceras and new eobeloceratids. MD III-D shows theentry of other new eobeloceratids and the earliest acantho-clymeniid, Pseudoprobeloceras. MD III-E shows the entry ofthe Petteroceratidae with Petteroceras, Meropharciceras, andthe acanthoclymeniid Ponticeras. By its close, all of the eo-beloceratids have become extinct as have pharciceratids apartfrom Neopharciceras and one species of Petteroceras.

Frasnian

Manticoceras Stufe, UD I (A–L).–This division correspondsrather exactly with the Frasnian as now defined. It corre-sponds to the dominance of the Gephuroceratidae and Be-loceratidae. Formerly it was divided into I beta, I gamma,and I delta by Wedekind (1913, 1917, 1918). Matern(1931a) combined the two middle zones as I beta (gamma).But work on far more detailed sections than were availableto those authors has led to the new subdivision into 12 di-visions, UD I A-L, the distinction of which is based on workin Australia (Becker et al., 1993), Russia (Becker et al., 2000;House, Menner et al., 2000), North Africa (Becker &House, 2000a, b), Germany (House & Ziegler, 1977;Becker &House, 1993, 1994a), France (House et al., 1985;House, Becker et al., 2000; House, Feist & Korn, 2000),and North America (House, 1962; House & Pedder, 1963;Kirchgasser, 1974, 1975, 1982, 1985; Kirchgasser &House,1981; House & Kirchgasser, 1993, and herein).

The fauna of UD I-A, or the PonticerasGenozone or Di-vision, is subdivisible into three phases, not all of which arerecognized internationally. At the very base, as shown in the


basal Frasnian GSSP (Global Boundary Stratotype Sectionsand Points) at Puech de la Suque, Montagne Noire, south-ern France (Klapper et al., 1987; House, Feist & Korn,2000), the record indicates that Peterroceras and Neopharci-ceras, the last remnants of the Pharciceratidae, just cross theboundary; their extinction early in UD I-A marks theFrasnes Event (House, 1985). Then the long-ranging Pon-ticeras and Epitornoceras continue with the entry of Chuto-ceras marking the base of an upper phase. UD I-B, or theKoenenites Genozone, is marked by the entry of Koenenitesand the Koenenitidae. UD I-C, the Timanites Genozone,corresponds to the curious international occurrence of thatgenus in the Timan Basin, Russia, western Canada, andWestern Australia (but not New York) and of the entry ofManticoceras perhaps a little higher as shown in WesternAustralia. UD I-D, the Sandbergeroceras Genozone, corre-sponds to the first entry of the name genus and to the Mid-dlesex Event.

UD I-E, the Probeloceras Genozone, corresponds to theentry of the name genus and of Gogoceras and Uchtites. UDI-F, the Prochorites Genozone, corresponds to the entry ofthat genus and probably a number of gephuroceratid generasuch as Carinoceras, Maternoceras, and Sphaeromanticoceras.UD I-G, the Mesobeloceras Genozone, corresponds to theentry first of Naplesites, then its descendant Mesobeloceras,and gephuroceratids such as Serramanticoceras and Costa-manticoceras. The base of UD I-H, the Beloceras Genozone,corresponds to the point of derivation of the name genusfrom Mesobeloceras; in restricted environmentsWellsites andSchindewolfoceras are thought to belong here but improvedcorrelation is desirable. UD I-I, the Playfordites Genozone,corrsponds to the entry of the name genus and ofTimanoceras, Trimanticoceras, Virginoceras, and the pseudo-ceratiteDevonopronorites (dated by R. T. Becker on evidencefrom Iran). UD I-J, the Neomanticoceras Genozone, corre-sponds to the entry of the name genus and of Clauseniceras.UD I-K, the ArchocerasGenozone, corresponds to the spreadof the index genus with the Lower Kellwasser Event andDel-phiceras; very early Crickites can occur here. UD I-L, theCrickites Genozone, corresponds to the acme of the namegenus and the range of Crickites holzapfeli (Wedekind,1913), the zone fossil of Wedekind’s I delta; Enseites andCeratobeloceras are limited to this level. All Gephurocerati-dae, Acanthoclymeniidae, and Beloceratidae are extinct bythe close of UD I-L, associated with the last stage of theUpper Kellwasser Event.

FamennianFollowingWedekind (1917, 1918), the Famennian has beendivided into ammonoid divisions II-VI, a practice con-firmed by Schindewolf (1923) and others. Originally these

divisions corresponded to the local German Famennianstage-type divisions. Thus, Division II, the Cheiloceras Stufe,equated with the Nehden; Division III, the Prolobites Stufe,and Division IV, the Platyclymenia Stufe, equated with theHemberg; Division V, the Clymenia Stufe, equated with theDasberg; and Division VI, the Wocklemeria Stufe, equatedwith theWocklum. But these stage names are in the processof being redefined in Germany and it is to be expected thatthis simple scheme, with its priority, is likely to be replaced.The terms were originally lithostratigraphic ones, and con-version to biostratigraphic terms, or chronostratigraphic ifaccepted by the International Union of Geological Sciences(IUGS), will require definition by GSSPs. Schindewolf(1934) combined Divisions III and IV as the Prolobites-Platyclymenia Stufe. The Roman-numeraled divisions ofWedekind were retained in the classification developed byBecker (1993a, b), which is now subdivided using theRoman alphabet, rather that the Greek as used byWedekindand his followers (Becker & House, 2000).

Cheiloceras Stufe, UD II, Nehdenian.–Now divided intonine divisions (UD II-A through I), of which the first six(II-A through F) correspond toWedekind’s II alpha, and re-mainder (UD II-G through I) corresponds to the former IIbeta, initiated by the entry of sporadoceratids. The Cheilo-ceras Stufe equates with the Nehdener Schichten or Nehden-ian according to the original definitions of Paeckelmann(1924).

The fauna of UD II-A, the Phoenixites Genozone, is animpoverished one following the Upper Kellwasser extinc-tions and is characterized by the opportunistic spread ofPhoenixites, a genus first recorded early in the Frasnian (UDI-C). UD II-B, the Cheiloceras (Raymondiceras) Genozone, ischaracterized by the entry of simple-suture cheiloceratidswith several species of Falcitornoceras. UD II-C, the Cheilo-ceras (Cheiloceras) Genozone, shows the entry of the trueCheiloceras and the tornoceratids Polonoceras and Oxy-tornoceras. UD II-D, the Paratorleyoceras Genozone, showsthe entry of advanced cheiloceratids with pointed adventi-tious lobes (A), and includes Cheiloceras (Staffites), Parator-leyoceras, Torleyoceras, and the tornoceratid Armatites. UDII-E, the Praemeroceras Genozone, shows the entry of thename genus. UD II-F, the Paratornoceras Genozone, showsthe entry of oxyconic dimeroceratids such as Paratornoceras.UD II-G, theMaenecerasGenozone, shows the entry of spo-radoceratids, at first with only shallow A2 lobes. UD II-H,the PosttornocerasGenozone, marks the entry of the true Spo-radoceras with deep A2 lobes and of the name genus. UD II-I, theDimerocerasGenozone, marks an international spreadof this genus (which appears rarely in UD II-H).



Prolobites Stufe, UD III (with UD IV forming the Hem-bergian).–This is divided into three divisions. UD III-A, thePernoceras Genozone, corresponds to the entry of thetornoceratids Pernoceras, Protornoceras, and Tornia. UD III-B, the Pseudoclymenia Genozone, is dominated by thePseudoclymeniidae. UD III-C, the Prolobites Genozone, ischaracterized by the entry of many groups of clymeniids.

Platyclymenia Stufe, UD IV.–This is divided into three di-visions, UD IV A-C, based on the entry of various clymenidgenera. UD IV-A, the PrionocerasGenozone, is characterizedby the Platyclymenia annulata (Münster, 1832) group, andmarked by the entry of prionoceratids. UD IV-B, the Pro-toxyclymenia Genozone, is characterized by the name genuswith early Cymaclymenia and Uraloclymenia. UD IV-C, thePachyclymeniaGenozone, is marked by the entry of Protoxy-clymenia (Franconiclymenia) and probably the earliest Platy-clymenia (Spinoclymenia), Pachyclymenia, and Costaclymenia.

Clymenia Stufe, UD V, Dasbergian.–This is divided intothree divisions, UD V A-C, based on the entry of variousclymenid genera. UDV-A, the ClymeniaGenozone, is char-acterized by C. laevigata (Münster, 1832) and the spread ofDiscoclymenia, Kosmoclymenia, Sphenoclymenia, Kiaclymenia,Biloclymenia, and others. UD V-B, Ornatoclymenia Geno-zone, is marked by the entry of O. ornata (Münster, 1834),with Gonioclymenia, Cycloclymenia, and distinctive speciesof Kosmoclymenia. UD V-C, the Kalloclymenia Genozone,is indicated by the entry of Piriclymenia piriformis (Schmidt,1924), and of Kalloclymenia.

Wocklumeria Stufe, UD VI, Wocklumian.–This is dividedinto six divisions, UD VI A-F, based on the entry of variousclymenid genera. UD VI-A, the Linguaclymenia Genozone,is marked by the entry of Kosmoclymenia (Muessenbiaergia)bisulcata (Münster, 1840), Sphenoclymenia brevispina (Lang,1929), and Linguaclymenia, and corresponds to the Lowersubarmata Zone and lower half of the Upper subarmataZone of Korn (1986). UD VI-B, the Balvia Genozone, ismarked by the entry of the name genus and of variousspecies ofGlatziella and was considered an upper part of theUpper subarmata Zone by Luppold et al. (1994). UDVI-C,the Parawocklumeria Genozone, is marked by the entry ofthe name genus and of Kamptoclymenia and Triaclymenia;this is the Lower paradoxa Zone of Korn (1986). UDVI-D,the Wocklumeria Genozone, is indicated by the entry ofWocklumeria and of Synwocklumeria and Kielcensia. UDVI-E, the Cymaclymenia Genozone, marks the remnant of cly-meniids after the extinctions of the Hangenberg Event. UDVI-F, the Acutimitoceras Genozone, is marked by the entryof imitoceratids with evolute inner whorls, and Acutimito-

ceras (Stockumites) but still with Cymaclymenia (Korn, 1991);this straddles the Devonian/Carboniferous boundary of thenew definition (Paproth et al., 1991).

GONIATITE BIOSTRATIGRAPHY

OF THE LATE DEVONIAN IN NEW YORK

The ranges of late Devonian goniatites in New York plottedagainst the stratigraphic divisions and the alignments ofNew York Regional Zones against the international goniatitezonation are summarized in Text-figs 23-24. This updates aprevious review (House & Kirchgasser, 1993). For the EarlyandMiddle Devonian, not covered in this report, there havebeen summary reviews in the past (House, 1962, 1978,1981b; Kirchgasser, 1985; Woodrow et al., 1989). So far asa contribution to international detailed zonation is con-cerned, early work suffered from the problems associatedwith an understanding of the complex facies relationshipsof the Devonian of New York State. For the Middle Devon-ian, the elucidation of the Hamilton sedimentary cycles andtheir correlation in central and eastern New York by G. A.Cooper (1930, 1933) was a special landmark. Although theTully Limestone had been long recognized as an importantmarker (Cooper & Williams, 1935), it disappears throughintra-Devonian erosion west of the Finger Lakes (Text-fig.7). For the late Devonian, the classic work by Chadwick(1935b) clarified the long-standing miscorrelations betweenthe eastern clastic facies and the more western neritic andpelagic facies (Kirchgasser,1985). It might be true to say thatit was not until the fundamental review of Devonian corre-lations by Cooper et al. (1942) that a stratigraphical frame-work comparable with European sections was available. Bythen, Miller’s (1938) review of Devonian ammonoids hadbeen published, but that was a taxonomic, and not a bios-tratigraphic, treatment.

Clarifications of the facies changes since 1950 have beenfundamental to this work, especially the two correlationcharts of L. V. Rickard (1964, 1975), which incorporatedthe results of correlations in the western New York Frasnianat a very fine level. Also important has been the work of Sut-ton (1960, 1963), Sutton et al. (1962), Colton & de Witt(1958), deWitt & Colton (1959, 1978), Pepper & deWitt(1950, 1951), and Pepper, de Witt & Colton (1956). Ofmajor significance to paleoenvironmental and sedimento-logical interpretation have been the many contributions byG. Baird and C. Brett (e. g., Brett & Baird, 1996).

An attempt at correlation of the New York sequence withEurope was made by House (1962). So far as an attempt ata local (New York) zonation is concerned, a start was madeby House (1968) and continued in a review of the am-monoids of the Appalachians (House, 1978). In the latterwork, numbers were given for faunas, and these are shown

Text-fig. 23. Ranges of late Devonian goniatites in New York State plotted against the stratigraphic divisions. Updated fromHouse & Kirchgasser (1993). Near top, read Pharciceras? sp.


Text-fig. 24. Alignment of conodont zones (Standard and Montagne Noire) and goniatite zones and divisions (after Becker & House,2000), with New York rock units. At the top, read Oswayo (for Osweyo).



in Text-figs 23-24. Since that time, international work bythe authors, usually jointly with R. Thomas Becker of theUniversity of Münster, Germany, has led to a much moredetailed understanding of international faunas. As far asNew York is concerned, our work in Western Australia(Becker et al., 1993) is particularly important.

REGIONAL NEW YORK GONIATITE ZONES

Zonation schemes are transitory in the sense that they aresubject to improvements and changes as new evidence be-comes available. This section updates zonations publishedearlier (House, 1978; Kirchgasser & House, 1981; House& Kirchgasser, 1993) and explains the correlation adoptedwith the international scheme of Becker & House (2000).This monograph covers the Upper Devonian in the classsicalsense, that is, starting with the Tully Limestone. The newGSSP redefinition of the base of the Frasnian Stage andUpper Devonian Series (Klapper et al., 1987; House, Feist& Korn, 2000) has led to arguments on the taxonomy ofconodont species thought to define the boundary (Johnson,1989; Ziegler & Sandberg 1994, 1996; Sandberg et al.,1988; Racki & Wrzolek, 1989; Racki & Bultynck 1993;Klapper, 1988, 1989, 2000); some of this history has beenreviewed by House, Feist & Korn (2000). But, under therules of the International Commission on Stratigraphy(ICS), the boundary is defined by the agreed point in theGSSP at Puech de la Suque in southern France and not byconodonts or goniatites. Some new regional zones are inte-grated into the numerical system of House (1978) andKirchgasser & House (1981) but further subdivided in thelight of new evidence. Text-fig. 24 shows the New York Re-gional Zones correlated with the conodont zonation schemeof Klapper (1989, 1997), following Becker &House (2000:fig. 1, table 4); see also Klapper & Becker (1999) and House(2002).

Following the methodology of House & Kirchgasser(1993) and Becker &House (2000), the Regional New YorkGoniatite Zones are essentially defined by the first occur-rence of a single species or, in some cases, a group of speciesor fauna.

GivetianThe Givetian Stage probably commences in New York a lit-tle above the Cherry Valley Limestone and is indicated ingoniatite terms by the true tornoceratids (House, 1981b)found in the Chittenango black shale of Cooper et al. (1942)and Rickard (1975). This could be a local representation ofthe Kacák Event (House, 1985, et seq.; 2002) .

Maenioceras Stufe, MD II (Faunas 7-12).–Several goniatitefaunas have been found in the beds up to the Tully Lime-

stone (House, 1978: faunas 7-12) and reviews have beenpublished (House, 1962, 1981b). New work by G. Kloc isnot published; perhaps most significant is his discovery ofMaenioceras for the first time in New York, as the guide tothe Maenioceras Stufe and MD II. Agoniatites occurs and isunder study by S. Klovac (American Museum of NaturalHistory, New York), and tornoceratids are abundant andwere monographed by House (1965). The distinction oftornoceratids from below and above the Centerfield is im-portant because it seems that only late Hamilton tornocer-atids can be derived in the remanié and pyritic lags such asthe Leicester Pyrite. Huddle (1981) demonstrated the post-Tully conodont ages of the pyrite lenses that also contain aderived goniatite fauna (Fauna 12) west of the last Tully out-crops (Text-fig. 7).

When the tornoceratid study was done, the HungryHollow Formation of Ontario was correlated with the Cen-terfield Limestone of New York, essentially followingCooper et al., (1942). Just above the Centerfield, at Moon-shine Falls, Cayuga Lake, Polygnathus timorensis Klapper,Philip & Jackson, 1970, is recorded, which indicates theconodont Lower varcus Zone (Klapper et al., 1970; Klapper,1981). Thus occurrences of Tornoceras uniangulare arkonenseHouse, 1965, in the Ontario Arkona Shale (below the Hun-gry Hollow) and Tornoceras uniangulare widderi House,1965 (Pl. 26, fig. 5) in the OntarioWidder Shale (above theHungry Hollow) were thought to correlate to positionsbelow and above the Centerfield Limestone in New York.Subsequently Brett & Baird (1985) questioned the HungryHollow-Centerfield correlation, interpreting the Centerfieldas a regressive unit within deeper water sediments and theHungry Hollow as the basal unit of a transgressive sequence.Landing & Brett (1987) discovered a regional disconformitybeneath the Hungry Hollow, but they interpreted the upperpart of the Hungry Hollow and lower Widder Shale as anofflap-onlap cycle within the P. timorensis Zone and thuscomparable to the upper part of the Centerfield. This issuewas taken up by Sparling (1992) who found a fauna includ-ing P. timorensis and P. ansatus Ziegler & Klapper, 1976, (theguide to the Middle varcus Zone) in the Prout Limestone ofErie County, Ohio. Because the Prout has generally beenconsidered of the same age as the Hungry Hollow, Sparlingused this presumption to argue that the Hungry Hollow isalso of Middle varcus Zone age. But the Prout fauna is rich,and that of the Moonshine Falls locality is poor, and it couldbe that the latter might still yield P. ansatus. More recentlySparling (1999) has argued that, because the Tully has a lateMiddle varcus and younger conodont fauna, the HungryHollow should be correlated with the Tully Limestone. Allof this neglects the clear fact that the varcus Zone spans con-siderable time (House, 1995), and the Middle varcus Zone

could span from a level near the Centerfield to the earlyUpper Tully (Text-fig. 7). As will be discussed later, the de-rived goniatite faunas of the Leicester Pyrite are only of thepost-Hungry Hollow type. Thus Sparling’s correlation of theWidder Shale and Hungry Hollow with theTully Limestoneis not accepted, although all could fall within part of theMiddle varcus Zone (see Aboussalam& Becker, 2001: fig. 1;Aboussalam, 2003: fig. 2).

Pharciceras Stufe.–This division was introduced by House(1985) for that part of the former Upper Devonian Zone ofPharciceras lunulicosta (G. & F. Sandberger, 1850) (I alphain Germany) that was transferred to the Middle Devonianwith the redefinition of the base of the Upper Devonian(Klapper et al., 1987) by the International Commission onStratigraphy (ICS) and International Union of GeologicalSciences (IUGS). Following work in Morocco, Becker &House (1994b) have established five zonal subdivisions(MD III A-E) as described earlier.

Regional Zone of Pharciceras amplexum (Fauna 13, MDIII-A).–The zonal species is only known from a single levelin the Upper Tully Limestone (Moravia Bed or Bed G of theWest Brook Member) as recounted earlier (see also House,1962, 1965; Kirchgasser & House, 1981) (Text-fig. 6). Onthe eastern side of Seneca Lake, where the Tully is well de-veloped, P. amplexum occurs 0.38 m above the Bellona CoralBed in Lodi Glen (Mill Creek, Seneca County; Loc. 3). AtJune’s Quarry, Tully (Onondaga County; Loc. 7), the levelis 1.27 m above the Coral Bed. The genus occurs at severalmore eastern localities as far as New Lisbon (OtsegoCounty). See details under Tully Formation in the localitysection.

Regional Zone of Pharciceras sp. (Fauna 14, MD III-?A).–The records of Pharciceras sp. from the Fillmore GlenBed of the uppermost Tully at Sheldrake and Mack creeksand other localities (Loc. Gen-1) clearly postdate the P. am-plexum horizon, but because none of the specimens knownshow sutures, they cannot be assigned.

Regional Zone of Ponticeras perlatum (Fauna 15a-c, MDIII D-E/UD I-A).–There appear to be three local phases ofthis fauna. Fauna 15a is marked by the entry of Epitornoceras(E. cf.mithracoides) in the upper Geneseo Shale, 12.2 m (40ft) below the Lodi Limestone, in Lodi Glen (Loc. 27), andapproximately 24-27 m (80-90 ft) above the Tully Lime-stone; the genus ranges to the middle Penn Yan Shale andequivalents.

Fauna 15b is marked by the entry of Ponticeras perlatumin the upper Geneseo Shale, immediately below the LodiLimestone (House, 1962), for example in Hubbard’s Quarry(Loc. 27b); the Lodi Limestone yields the best preserved ma-terial (Kirchgasser, 1975). The conodont norrisi Zone fauna

in the Lodi Limestone represents the latest Givetian (MiddleDevonian) in current definitions (Klapper et al., 1987), andthe base of the Frasnian and Upper Devonian is taken wherethe Lower asymmetricus Zone (MN1) (or falsiovalis Zone)occurs at the junction of the Lodi Limestone and the over-lying Penn Yan Shale (Kirchgasser, 1994).

FrasnianManticoceras Stufe.–Because neither Neopharciceras or thecritical species of Petteroceras are known in New York, it isconodont evidence that places the Givetian/Frasnian bound-ary precisely and it falls within our Fauna 15b.

Fauna 15c is marked by the entry of Chutoceras [C.nundaium, formerly Ponticeras cf. regale (Holzapfel, 1899)]in the Ithaca Shale and Sandstone, at approximately the levelof theWilliams Brook Coquinite, approximately 53 m (175ft) above the Renwick Shale at Williams Brook (Loc. R, I-2) and 72 m (235 ft) above the base of the Renwick Shalein Fall Creek, Ithaca (Loc. X, Dy-10); ?C. nundaium occursat Cascadilla Creek (Loc. Z) and perhaps Enfield Glen (Loc.BB). These Ithaca Formation occurrences of Chutoceras nearCayuga Lake are equivalent in the west to the interval in thePenn Yan Shale between the Lodi Limestone and the CrosbySandstone (Kirchgasser, 1985). These records are the mostprecise evidence internationally of the entry of Chutoceras.

Regional Zone of Koenenites styliophilus styliophilus(Fauna 16a–b, UD I-B).–Koenenites enters in the upperPenn Yan Shale at the Linden Goniatite Horizon (Kirch-gasser & House, 1981) at Linden (Loc. 15) and is foundeastward to Canandaigua Lake. It occurs with Tornoceras ar-cuatum in the Crosby Sandstone (which is probably slightlyyounger than the Linden Horizon), near Keuka Lake. Theentry of the genus defines the base of UD I-B. The genus islast seen (as defined here) in the West River Shale. Twophases seem recognizable:

Fauna 16a, the earliest phase, is dominated by Koenenitesstyliophilus styliophilus, which is restricted to the Upper PennYan Shale and is best known from the Linden Horizon andCrosby Sandstone and associated styliolinid beds. Acantho-clymenia (formerly identified as Probeloceras) enters in thisphase.

Fauna 16b is characterized by Koenenites styliophilus kil-foylei n. ssp. of the lower part of the Genundewa Limestone,especially at Bethany Center (Loc. 24/1) with Acanthocly-menia genundewa and Tornoceras uniangulare compressum.Koenenites? fasciculatus occurs at the type area of the Genun-dewa Limestone at Canandaigua Lake (Loc. 23b). K.? fasci-culatus is also known from the Genundewa Limestoneequivalents near Keuka Lake but with conforming materialfrom the earliest West River Shale (Locs DD, FF).

Regional Zone of Manticoceras contractum (Fauna 17a,


UD I-C).–Manticoceras enters in the upper GenundewaLimestone at Geneseo (Livingston County; Locs 19 and19a) and subsequently the genus is not seen until the upperWest River Shale. Together with M. contractum, two otherspecies were recorded from the Genundewa by Clarke(1898):M. nodifer andM. apprimatum; Acanthoclymenia cf.genundewa also occurs. This is the level that was formerlytaken to indicate the base of the M. nodulosum (Wedekind,1913) (I beta) Zone (House, 1962). The entry of Mantico-ceras represents a transgressive pulse. It is to be expected thatTimanites should precede this entry but there is no evidenceof this form in the Appalachian Basin; that name is preferredfor the division UD I-C, the Timanites Genozone, becauseit is a short-lived genus whereasManticoceras continues untilthe close of the Frasnian. In any case, the Genozone (or Di-vision) terminology is one resulting from international,rather than local, work. There is a gap in the record in thesucceeding lower West River Shale. The type species forManticoceras, M. simulator (Hall, 1874), was recorded asfrom the “Leiorhynchus Beds” of the Naples Formation at ornear Ithaca, New York (see holotype and only known spec-imen, Pl. 9, Figs 4-5). The precise horizon in the IthacaShale and Sandstone is not known but it is expected to be inthe equivalents of the West River Shale, and probably highrather than low in that unit (Kirchgasser, 1985).

Regional Zone of Koenenites beckeri n. sp. (Fauna 17b,UD I-C).–K. beckeri n. sp., formerly referred to K. aff. lamel-losus (G. & F. Sandberger, 1851), occurs in the West RiverShale, immediately below the Bluff Point Siltstone. Onespecimen is close to the specimen described from the Mon-tagne Noire under the name Hoeninghausia aff. archiaciGürich, 1896 (House et al., 1985: text-figs 8D-E, pl. 3, figs10-12), but we decided that it is better to keep the genusHoeninghausia for undoubtedly oxyconic forms. The upperWest River Shale from the Bluff Point Siltstone to the top ofthe unit has horizons that yield koenenitids of variable qual-ity of preservation and size, hence assignments are corre-spondingly difficult. Records include Acanthoclymenia aff.neapolitana, Lobotornoceras aff. hassoni, and Manticoceraslamed cf. cordatum. There is an evolute form that has beenreferred to as “Archoceras“ but that we now assign to Pon-ticeras? sp.

Regional Zone of Sandbergeroceras syngonum (Fauna 18,MD I-D).–The zonal species is the only form known fromthe Middlesex Shale and it has been widely reported fromNew York and the Appalachians. S. syngonum was describedfrom Snyder’s Gully (Loc. 46c), Canandaigua Lake. Thismultilobed and ribbed form is assigned to the Triainocerati-dae and there is a long gap in the record of this group backto its presumed origin in the late Givetian. As with someother multilobed genera, its occurrence seems related to a

hypoxic transgressive phase. We have not searched the Mid-dlesex Shale systematically; it would appear likely that thisgenus is probably confined to a narrow band within theMiddlesex, perhaps at the time of maximum transgression.We have found specimens of Sandbergeroceras in Middlesexequivalents in Tioga County (Loc. 64).

Regional Zone of Probeloceras lutheri (Fauna 19, UD I-E).–Typical faunas of the former Manticoceras cordatumZone (I gamma) enter with the Cashaqua Shale in New YorkalthoughM. lamed aff. cordatum occurs earlier in the upperWest River Shale. In the lower Cashaqua Shale, P. lutheri en-ters and is common in the middle part and up to, but notincluding, the Shurtleff Septarian Horizon.M. sinuosum sin-uosum occurs throughout the Cashaqua Shale andM. s. tar-dum and M. s. clausium n. ssp. are restricted to intervals inits lower part. Aulatornoceras eifliense is present in the middleof the Cashaqua Shale.

Regional Zone of Prochorites alveolatus (Fauna 20, UD I-F).–In the upper Cashaqua Shale, a new fauna appears thatis characterized by P. alveolatus, a senior synonym of P. strixKirchgasser, 1968. Its entry is with the celebrated bariticfauna of the Shurtleff Septarian Horizon, known best fromShurtleff ’s Gully, Livonia (Livingston County; Loc. 41),which yielded the type material of Acanthoclymenia neapoli-tana (see House, 1961) but this form already appears slightlyearlier. Manticoceras sinuosum sinuosum and Tornoceras uni-angulare obesum are abundant in the Shurtleff SeptarianHorizon, and Aulatornoceras eifliense is also present.

Regional Zone of Naplesites iynx (Fauna 21a, UD I-G).–The horizon(s) of the type material forN. iynx (Clarke,1898) and N. naplesense (Clarke, 1898) is not known butthe specimens are said to have come from the Naples area ofOntario and Yates counties. In view of the careful searchingof the Cashaqua Shale, we feel that the horizon(s) must bein the lowermost Rhinestreet Shale. A major problem still re-maining is the need for careful searching and correlation oflevels yielding faunas in the Rhinestreet Shale, which is thethickest black shale of the Frasnian black shales in New Yorkand thought to represent a considerable period of time. Onthe international scale, it is known that Mesobeloceras (UDI-G) is a later development of Naplesites but, as now inter-preted, that form is not known in New York and neither isthe descendant Beloceras (which enters in UD I-H).

Regional Zone of Wellsites tynani (Fauna 21b, UD-?H).–This division approximately corresponds with the Be-locerasGenozone but we do not believe that true Beloceras isrepresented in New York. In its place is a curious group ofmultilobed forms including Wellsites and Schindewolfoceras;these are found in the development of the Rhinestreet Shalein the central part of the state, especially south and southeastof Ithaca. Their occurrence well to the east and approaching


the more clastic facies of the Catskill Delta suggests that theyresult from particularly extensive transgressions within themajor transgression represented by the Rhinestreet Shale andcorrespond to the triainoceratids in the Middlesex Shale inrepresenting unusual deeper-water forms. The facies is verydifferent from the Beloceras limestones of Europe and Aus-tralia. The Rhinestreet examples raise problems of their exacttime equivalence with the western succession. Perhaps theearliest is the group represented by the genus Wellsites. W.tynani is from near Elmira (Chemung County; Loc. 70),and thought to be from the Moreland Shale, a basal tongueof the Rhinestreet Shale. Wellsites williamsi is from BaldMountain (Tioga County; Loc. 68), from a level probablyjust above the Moreland Shale.

Regional Zone of Schindewolfoceras chemungense (Fauna21c, UD I-?H-I).–This level marks the incoming of newforms with ribbed early whorls characteristic of the Triain-oceratidae. The source of the original specimen of S. che-mungense (Text-fig. 24I) was not precisely located but, whenit was refigured, Hall (1879: 69) gave the locality asChemung near Owego (Tioga County; Loc. 69). Otherspecimens referred to the genus with doubt are S.? equicosta-tum said to come from Athens, Pennsylvania, and later saidto come from a boulder there. The best localized specimenis S.? aff. equicostatum from Fairfield Forest (Loc. 67) nearSpeedsville (Tioga County) at a level between the MorelandShale and Roricks Glen Shale tongues of the RhinestreetShale, which Sutton &McGhee (1985) indicated to be stillrather low in the Rhinestreet but above the levels bearingWellsites.

Regional Zone of Playfordites cf. tripartitus (Fauna 22a,UD I-I).–There is a thick interval, mostly of black shale,within the Rhinestreet Shale that has yielded neither goni-atites nor conodonts. However, in the upper RhinestreetShale in sections near Lake Erie, small-scale rhythms withcalcareous levels or concretions occur and in this facies, andin equivalents in Gardeau Shale and Sandstone farther east,a suite of goniatite genera appear for the first time. At John-son Creek (Wyoming County; Loc. 58), in the concre-tionary horizon (Loc. 58/1) in the cycle below the cycle withthe Relyea Creek Horizon, a key marker species, Playforditescf. tripartitus (NYSM 12074) enters along with Carinocerasvagans,Manticoceras lamed, Sphaeromanticoceras rhynchosto-mum, and Linguatornoceras aff. linguum. The genus Play-fordites (type species: P. tripartitis) is the marker genus of UDI-I, the PlayforditesGenozone or Division. Playfordites is notknown to occur outside of its genozone (or division) inWestern Australia, where the genozone was delineated(Becker et al., 1993), or in Germany outside the classicManticoceras nodulosum Zone (I beta) of Wedekind (1913)or lower M. cordatum Zone of House & Ziegler (1977)

(Becker & House, 2000).The occurrence of Playfordites cf. tripartitus in the upper

Rhinestreet Shale in the bed at Johnson Creek (Loc. 58/1)defines the New York Regional Playfordites cf. tripartitusZone (22a), a zone that is restricted to that single level. Theconodont fauna of the Reylea Creek Horizon, in the over-lying cycle, based on collections to be described with G.Klapper, includes the marker species Palmatolepis semichato-vae Ovnatanova, 1976, which supports a correlation of theupper Rhinestreet P. cf. tripartitus occurrence with the con-odont MN Zone 11. In the standard conodont zonation,the level would be above the jamieae Zone, in the P. semicha-tovae interval of the Lower rehenana Zone (Klapper &Becker, 1999; Sandberg et al., 2002).

Playfordites cf. tripartitus (NYSM 12075) also occurs inthe lower Angola Shale, in the Point Breeze Goniatite Bedat Relyea Creek (Loc. 82/6), an anonomously high occur-rence that we are placing, for reasons given below, in theNeomanticoceras Genozone (UD I-J), that is, in the geno-zone above the Playfordites Genozone.

Regional Zone of Sphaeromanticoceras rhynchostomum(Fauna 22b, UD I-J).–The goniatite faunas in the intervalfrom upper Rhinestreet Relyea Creek Horizon to the top ofthe Angola Shale define the New York Regional S. rhynchos-tomum Zone (22b), named for the most common mantico-ceratid in the interval and particularly in the Relyea CreekHorizon of the upper Rhinestreet Shale and Point BreezeGoniatite Bed of the lower Angola shale. The placement ofthe S. rhynchostomum Zone in the Neomanticoceras Geno-zone or Division (UD I-J) and its differentiation from thepreceeding P. cf. tripartitus Zone (22a) is problematic: Thename genus for the Neomanticoceras Genozone has not yetbeen identified in New York, but already in the upperRhinestreet Shale occurs the similarly oxyconic genusCarinoceras, but that genus is internationally known to startin UD I-F, that is, much earlier. A search is required of theNew York oxyconic forms to see whether any develop theextra division in the ventral saddle that is characteristic ofNeomanticoceras.

In the cycle above the cycle with the Playfordites horizon(Loc. 58/1) discussed above, is the most productive concre-tionary level in the upper Rhinestreet Shale, the RelyeaCreek Horizon, named for a bed at Relyea Creek (Loc. 60/3)near South Warsaw (Wyoming County). This locality wasknown to Clarke (1899c) and possibly Luther (1903: 1012)under the name Gibson’s Glen. The fauna of the RelyeaCreek Horizon at Relyea Creek (Loc. 60/3) includesSphaeromanticoceras oxy, S. aff. oxy, Carinoceras vagans,Man-ticoceras lamed, and, in the bed above (Loc. 60/4), S. rhyn-chostomum and Linguatornoceras aff. linguum. Many of theseforms continue upward into the lower Angola Shale where


Aulatornoceras auris, A. aff. eifliense, and Carinoceras sororiumenter.

In the sixth cycle above the base of the Angola Shale is aparticularly rich level named the Point Breeze Goniaite Bed(Kirchgasser & House, 1981) after a locality on Lake ErieShore (Loc. 72/6). In addition to Playfordites cf. tripartitesfrom Loc. 82/6 discussed above, the Point Breeze GoniatiteBed and the bed in the cycle below at Hampton Brook (Loc.74/6a) contain Sphaeromanticoceras oxy, Carinoceras vagans,Manticoceras lamed, S. rhynchostomum, Linguatornoceras aff.linguum, A. aff. eifliense, andTornoceras cf. typum (which be-gins in Loc. 54/1 in the upper Rhinestreet), along with thefirst appearances of Aulatornoceras paucistriatum, Crasso-tornoceras aff. crassum, andM. aff. lamed. The higher beds inthe Angola Shale are poor in goniatites. M. lamed and C.aff. vagans occur in Loc. 79/10 at Varysburg and ?S. rhyn-chostomum occurs higher in the Angola above the “TrinityBlack” black shales at Loc. 18/14 near Varysburg (GassmanRoad).

The distinction between the Playfordites andNeomantic-oceras Genozones in New York is uncertain. Although Neo-manticoceras has not been found in New York, we haveplaced the New York Regional Sphaeromanticoceras rhynchos-tomum Zone (22b) of the upper Rhinestreet to the top of theAngola, in the Neomanticoceras Genozone or Division (UDI-J). In doing so, we assume that the occurrence of P. cf. tri-partitus in the Point Breeze Goniatite Bed of lower Angolashale represents an extension of the range of Playforditesabove its genozone into the succeedingNeomanticocerasDi-vision (UD I-I). Comparison of the S. rhynchostomum Zonefauna with the upper Manticoceras cordatum Zone (I delta)fauna is not particularly helpful because most key markerforms in Germany do not occur in New York, such as N.paradoxum (Matern, 1931a), Carinoceras galeatum(Wedekind, 1918), Trimanticoceras cinctum (Glenister,1958), Maternoceras sandbergeri (Wedekind. 1913) and rel-atives, and Clauseniceras expectatum (Wedekind, 1913).However, two forms in common, Crassotornoceras aff. cras-sum and Aulatornoceras paucistriatum, do occur in the PointBreeze Goniatite Bed and these support an assignment ofthe New York Regional S. rhynchostomum Zone to the Neo-manticoceras Genozone (UD I-J) (Becker & House, 2000:129-130). The conodont fauna of the Point Breeze Goni-atite Bed is not diagnostic but it could already be in MNZone 12. The poor goniatites of the upper Angola are as-signed to the S. rhynchostomum Zone.

Regional Zone not assigned (Fauna 23, UD I-K).–Nofauna is known from the Pipe Creek Shale that we considerto represent the Lower Kellwasserkalk level of Europe. Alsoin New York, there are no records of Archoceras, a genus thatin Europe becomes opportunistically abundant with the

Lower Kellwasser Event (Feist, 1990) although Archoceraswabashense (Kindle, 1901) is known from Indiana (House,1962) (Pl. 17, Fig. 1) along with Clauseniceras delphiense(Kindle, 1901) (Pl. 17, Figs 2-3). Detailed study has beenundertaken of the early Hanover faunas by G. Kloc, and weare indebted to him for some information, but his resultshave not been published. The distinction between UD I-Kand UD I-L in New York is not clear because the name gen-era are not known [but see a record of ?Archoceras from IrishGulf (Loc. 90) referred to below]; the assignment to thegenozones of Faunas 23 and 24 is therefore tentative.

Regional Zone of Delphiceras cataphractum (Fauna 23,UD I-?K).–The first occurrence of the name species D. cat-aphractum in the lowermost Hanover Shale defines the baseof the zone; it is a distinctive, small, rather evolute, and pe-riodically constricted manticoceratid that is not unlike Ar-choceras varicosum (Drevermann, 1901) but has a subdividedventral lobe. In the absence of Archoceras at this level in NewYork, a correlation with the ArchocerasGenozone or Division(UD I-K) is problematic.

Delphiceras cataphractum is common in the lowermostHanover Shale at Beaver Meadow Creek, Java (WyomingCounty; Loc. 92), where it occurs with Aulatornocerasrhysum in the first meter (Bed 92/1-2) of the Hanover Shale.AtWalnut Creek, at Silver Creek (Chautauqua County; Loc.89a), the first records of D. cataphractum are approximately1.8 m above the Pipe Creek Shale and they continue over 10m higher. The conodont age of the lowermost HanoverShale is uncertain. It is assigned here to MN Zone 12 (Klap-per et al., 1995) but to MN Zone 13 by Over (1997a, b),corresponding to the Lower or Upper rhenana Zone in thestandard zonation (Klapper & Becker, 1999).

Regional Zone of Crickites lindneri (Fauna 24a, UD I-L).–In the lower Hanover Shale at Walnut Creek (Loc. 89a)C. lindneri enters the section near Bed 89a/4 or 5; the firstoccurrence of this species defines the base of the New YorkRegional C. lindneri Zone and the International CrickitesGenozone or Division (UD I-L). At Walnut Creek, otherloose specimens of C. lindneri probably came from Bed89a/4 or 5. Bed 89a/5 is also the source of the specimen col-lected by G. Kloc that we earlier described as Sphaeroman-ticoceras aff. rickardi (House & Kirchgasser, 1993: 247, figs.G-I).

Regional Zone of Sphaeromanticoceras rickardi (Fauna24b, UD I-L).–The name form of this zone enters in thelower part of the middle of the Hanover Shale (a loose spec-imen from near Bed 91/1 at Glade Creek, north of Stryk-ersville, Wyoming County) and ranges to the upperHanover where a single specimen was found by L. Rickardin Bed 91/7 in Glade Creek, 7.6 m (25 ft) below the base ofthe Dunkirk Shale (Text-fig. 16). New York Faunas 24a and


24b together correlate with the lower subzone of the Crick-ites Genozone as defined by Becker & House (2000: 132).

Regional Zone Unnamed to top of Frasnian Stage(Fauna 24c, UD I-L).–Near the top of the Hanover Shale inBed 90/2 in the section at Irish Gulf, southeast of NorthBoston (Erie County), small goniatites with distinctivelyevolute and compressed inner whorls determined as ?Ar-choceras sp. (NYSM 16587 and 16588) occur with smallsubinvolute goniatites with a rounded, rather rectilinearwhorl section assigned as ?Crickites sp. juv. (Text-fig. 45H;NYSM 16589). The whorl section of ?C. sp. juv. is similarto that of C. holzapfeli, the highest Frasnian goniatite zone-fossil in Europe. The goniatites of Bed 90/2, constitutingFauna 24c, are the highest known goniatites in the NewYork Frasnian and they define an unnamed Regional Zone.Fauna 24c might correlate with the upper subzone of theCrickites Genozone as defined by Becker & House (2000:132).

Bed 90/2 is a 2.0-2.5 cm-thick baritic, crinoid and gas-tropod-rich, concretionary horizon that is 4.7 m below thebase of the Dunkirk Shale and 1.76 m below the conodont-defined Frasnian/Famennian boundary (Over, 1997a, b). Itis shown inText-fig. 16 (2 cm bed) and by a concretion sym-bol in Over’s (1997b) fig. 8. The conodont fauna of Bed90/2 indicates the highest Frasnian MNConodont Zone 13(Over, 1997b: fig. 8) and conodont linguiformis Zone of thestandard zonation (Klapper & Becker, 1999).

No higher Frasnian goniatites are known in New York,but possible goniatite anaptychi (ammonoid opercula) occurin the Huddle Rider Bed, a thin black shale a meter or sobelow the top of the Hanover in Glade Creek (Loc. 91, Text-fig. 16) and approximately 2 m below the Frasnian-Famenn-ian boundary. The conodont-defined Frasnian-Famennianboundary, as determined by Over (1997a, b) and shown inText-fig. 16, lies within the upper Hanover Shale or lowerDunkirk Shale (Locs 91 and 92) depending on where theHanover-Dunkirk boundary is drawn.

Homoctenids have been noted near the Hanover-Dunkirk boundary at some localities and these distinctivedacryoconarids were once thought to be extremely rare afterthe end-Frasnian [Schindler (1990) recorded only a singlespecimen above the Upper Kellwasserkalk], but now they areknown to range well into the Famennian, for example, inChina (Xing Li, 2000). At Beaver Meadow Creek (Loc. 92),homoctenids occur profusely in the first siltstone above theFrasnian-Famennian boundary (Over, 1997a, b; Over et al.,1997); this horizon is the first of the three stippled units inthe lower Dunkirk Shale shown by Pepper & deWitt (1951:Section 7, Java). Homoctenids are also abundant in blackshale 50 cm above the Frasnian-Famennian boundary inGlade Creek (Loc. 91).

Famennian

Cheiloceras Stufe.–Regional Zone of Cheiloceras (Cheiloceras) amblylobum

(Fauna 25, UD II-C).–The most significant level with thisfauna is the Corell’s Point Goniatite Bed of the GowandaShale, traceable from Lake Erie Shore at Corell’s Point (Loc.97), and generally eastward at several localities as far asMarshfield (Erie County; Loc. 106). Cheiloceratids referredto the name species continue eastward at least as far as Java(Wyoming County; Loc. 109) and probably farther to theeast. Fauna 25 comprises C. (C.) amblylobum, Truyolsocerasbicostatum, and Phoenixites concentricus. The name genusconfirms correlation with the Cheiloceras (C.) Genozone UDII-C (Becker & House, 2000).

Regional Zone ofTruyolsoceras clarkei [Fauna 26, UD II-?C (?II-B)].–There is an early level in the Gowanda Shalewith T. cf. bicostatum in Walnut Creek (Loc. 99a) nearForestville (Chautauqua County); the same village is men-tioned for the locality of the types of the very different formdescribed here as T. clarkei. Note that earlier (House 1962,1965), Miller’s species was referred to Aulatornoceras. Theexact level of the T. clarkei deposit has not been determinedbut it is not thought to be so high as to be described as nearthe Laona Siltstone. Further collecting is needed at these lev-els. The queried reference to Genozone UD II-B has to bequite tentative in view of the absence of simple-suturedcheiloceratids.

Regional Zone ofMaeneceras aff. acutolaterale (Fauna 27,UD II-G).–No representative of Oxytornoceras, Praemero-ceras, or Acrimeroceras, all guides to UD II-D-F, are knownin New York or North America. The next level, theMaeneceras Genozone UD II-G, is represented by a singlespecimen (USNM 137645) of M. aff. acutolaterale, previ-ously determined as M. cf. pompeckji, from the middle partof the Ellicott (Chadakoin) Shale near Summerdale (Chau-tauqua County; Loc. 110). R. T. Becker informed us thatthere are problems regarding the type specimen of M. pom-peckji, in which case that name would be replaced by M.acutolaterale. These forms with shallow A2 lobes have beeninvestigated by Bochwinkle et al. (n. d.), who referred thespecies to UD II-I, but the New York form is likely to beearlier.

Fauna 27 is the highest known in New York, but it isconvenient to carry the known succession of eastern NorthAmerica to the top of the Devonian by commenting on thefaunas of Pennsylvania and Ohio.

Regional Zone of Maeneceras milleri (Fauna 28, UD II-H).–The name form, a single specimen, has been describedby Flower & Caster (1935), Miller & Flower (1936), andMiller (1938). If Sporadoceras is defined by its pointed A2


lobes, rather than the depth of the A2 lobes, as proposed byBockwinkel et al. (1999), then Maeneceras is the correct as-signment. But the size at which the pointed A2 lobe isachieved is essential to the definition. The specimen is fromHoward Quarries (Erie County; Loc. 111; White, 1881:103), Pennsylvania, beside Falls Run, approximately 8 mi(12.8 km) from the Lake Erie Shore. The Lower Venango(Cattaraugus) Sandstone crops out just below the quarry.The Regional Zone of M. milleri correlates to the Post-tornoceras Genozone UD II-H of Becker & House (2000).

Prolobites and Platyclymenia Stufe.–Regional Zone of “Pleuroclymenia” ohioensis (Fauna 29,

UD IV-A).–No faunas equivalent to the Prolobites Stufe areknown, unless ?Genus edwinhalli (Clarke, 1898), a speciesthat in this report is unassigned, is included. The type local-ity for the P. ohioensis Zone level is on theWest Branch of theVermilion River, 550 yd (490 m) northwest of the NY Rte.113 road bridge over the river at Birmingham, 12 mi (20km) west of Cleveland, Ohio. The fauna comes from theupper part of the lowest 1.8 m of the Cleveland Shale belowa siltstone unit and includes Platyclymenia spp., P. ohioensisHouse, Gordon & Hlavin, 1986, Cyrtoclymenia, andMaeneceras inflexum (Wedekind, 1908) (House et al., 1986).Fauna 29 appears to be early in the Platyclymenia Stufe, orPrionoceras Genozone UD IV-A (Becker & House, 2000).

Clymenia and Wocklumeria Stufe.–Regional Zone of Cymaclymenia sp. (Fauna 30, UD VI-

E).–No goniatite faunas equivalent to the Clymenia Stufe(UD V) are known in eastern North America. There is alevel near Rockford, Indiana, which has yielded a specimenreferred to Epiwocklumeria? sp. that might correspond to theWocklumeriaGenozone UDVI-D (Becker &House, 2000)but that cannot yet be placed in the Ohio succession (Houseet al., 1986).

From 0.6-1.5 m (2-5 ft) below the top of the ClevelandShale on Chance Creek (Lorain County), Ohio, and someother localities, Cymaclymenia sp. occurs as flattened pyriticfilms together with poor indeterminable sporadoceratids andprionoceratids (House et al., 1986). Fauna 30 correlates tothe Wocklumeria Stufe, Cymaclymenia Genozone UD VI-E(Becker & House, 2000).

Regional Zone of Rectimitoceras quadripartitum (Fauna31, UD VI-?F).–Faunas near the base of the Bedford Shale,for example, near Bedford (Cuyahoga County), Ohio, 1.7mi (2.7 km) northeast of Amherst City Hall, and TinkersCreek and neighboring localities (House et al., 1986), yielda fauna including R. (Prionoceras) quadripartitum (Münster,1939) without any other definitive Famennian forms. Thefauna is thought to be high in theWocklumeria Stufe, per-

haps UD VI-F.

SUMMARY

Thus there is represented in New York and adjacent states anexcellent succession of late Devonian ammonoid faunas. Be-cause of the thick nature of the succession, stratigraphicalprecision is as fine as anywhere in the world. Only a few lev-els are rich, however, and those are mostly in the Frasnian.In general, the faunas are of low diversity and abundanceand this is thought to be because of the restricted environ-ment provided by the Appalachian embayment in whichthey occurred, an environment that probably did not haveregular access to pelagic environments in which most De-vonian ammonoids flourished, as was long ago noted byClarke (1902). The individual horizons at which faunasoccur, however, might well represent the occasions whensmall rises of sea level enabled ammonoid faunas to enterthe basin, and in the future these marine bands might serve,as in the Carboniferous, as ecological as well as chronostrati-graphic markers.

Some discussion of the transgressive pulses that seem rec-ognizable in the New York succession and to which the entryof new goniatite faunas, or marker bands, seem related hasbeen documented elsewhere (House 1983; House & Kirch-gasser 1993; House et al., 2000) and has been also covered,especially with relevance to dating of events by conodonts,by Johnson et al. (1985, 1986), Klapper et al. (1995), andKlapper & Becker (1999).

The faunas reported here are from a considerable areaand from a substantial thickness of strata. Clearly, we havenot exhausted knowledge of the ammonoids of this classicand beautiful area, but we hope the time framework laiddown in this work will enable others to fill in the many miss-ing gaps that doubtless remain.

SYSTEMATIC PALEONTOLOGY

INTRODUCTION

In the taxonomic descriptions that follow, lengthy syn-onomies are not included and concentration is made on thedescription of type material. Within the families, the nom-inate genus is described first and other genera are treated inapproximate stratigraphical order. The same pattern is fol-lowed within genera as far as possible. Abbreviations of mor-phological terms used are as given in Text-fig. 25 or areexplained as they are used. The general terminology for shellform, sutures, and growth lines has changed little since theaccounts of Miller (1938) and Moore (1957), but more de-tailed accounts of sutural terminology are available (Schin-dewolf, 1954; Kullmann & Wiedmann, 1970; Wiedmann& Kullmann, 1981; House, 1996).


The section Distribution gives the stratigraphic and ge-ographic distribution of each taxon. Under the headingRange, the biostratigraphic position of each taxon is listed inthe following order: (1) name of Stage (e. g., Givetian), (2)genus name and key of International Genozone or Division[e. g., Pharciceras Genozone MD (Middle Devonian) III-A]following Becker & House (2000), (3) name of New YorkRegional Zone and Fauna number in Appalachian Basinsuccession [e. g., Pharciceras amplexum (13)] followingKirchgasser & House (1981), House & Kirchgasser (1993),and Becker & House (2000), and (4) name of StandardConodont Zone (e. g., Middle varcus Zone) following Sand-berg et al. (1989), Ziegler & Sandberg (1990) and Klapper& Johnson (1990), and, for the Frasnian Stage, Zone Num-ber of Montagne Noire (MN) Conodont Zone followingKlapper (1989, 1997), Klapper et al. (1995), and Klapper &Becker (1999).

REPOSITORIES

The major collections referred to, including the bulk of Halland Clarke’s material from the nineteenth century, are in theNew York State Museum, Albany (NYSM), which is wheremost of our own collections have been deposited. Some earlymaterial is in the American Museum of Natural History inNew York (AMNH). The large collection, mainly by H. S.Williams and especially J. W. Wells, formerly at CornellUniversity Museum of Paleontology (CU), is now either in

the NYSM or the Paleontological Research Institution(PRI), Ithaca, where the Cornell collection retains its origi-nal numbering. The PRI also holds much other material.Some early material of H. S.Williams and material collectedby G. A. Cooper is in the U. S. National Museum (USNM),Washington, DC (now National Museum of Natural His-tory), with some material collected more recently. The avail-able material collected by E. M. Kindle is held in theGeological Survey of Canada, Ottawa (GSC). Some speci-mens referred to are in the Buffalo Museum of Science(BMS), the geological collections at the University of Michi-gan, Ann Arbor (AAM), the Deparment of Geoscience atthe University of Iowa, Iowa City (SUI), the Sedgwick Mu-seum, Cambridge (SM), the University Museum, Oxford(OUM), the University ofWestern Australia, Perth (UWA),the Musée royale d’Histoire Naturelle, Brussels (IG), theMuseum für Naturkunde der Humboldt Universität, Berlin(HUM), and the Museum für Naturkunde, Berlin (HMMBC).

The material resulting from our earlier work (House,1962, 1965; Kirchghasser, 1975) has already been depositedin the NYSM and cataloged (Kilfoyle, 1969). The bulk ofthe material collected after 1965 is deposited in the NYSM;this material was initially labeled in green paint with blacklocality numbers in the 3000s and each specimen in an in-dividual collection numbered (e. g., 3136/9). A catalog ofthese numbers has been deposited with the NYSM. Speci-mens figured, or referred to, have been assigned NYSMnumbers. Other fossil material is deposited in the NYSMstratigraphical collection. Copies of our field notes, sectionlogs and catalogs are also deposited in the NYSM.

PhylumMOLLUSCA Cuvier, 1897Class CEPHALOPODA Leach, 1817

Subclass AMMONOIDEA Agassiz, 1847Order ANARCESTIDAMiller & Furnish, 1954Superfamily PHARCICERATOIDEA Hyatt, 1900

This superfamily is the first of the two major stocks showinga progressive proliferation of umbilical lobes on the flanks;the second is the Gephuroceratoidea. Both characterize, orrather, define, the German Adorfian as originally used(Wedekind, 1913). The families included in the Pharcicer-atoidea are the Pharciceratidae, Petteroceratidae, Eobelocer-atidae, Devonopronoritidae, andTriainoceratidae. Only thefirst and last are represented in New York State.

The Pharciceratidae and Triainoceratidae are well-de-fined groups that are discussed in further detail below. Bothshow a morphological series of forms of successive suturalcomplexity, the distinction between them resting on the co-station and nodation of the latter. The Eobeloceratidae is alate Givetian group that does not show the typical pinchedText-fig. 25. Diagrams illustrating terminology used in goniatite

descriptions and measurements given in the text.



lateral lobes of the Pharciceratidae. The Givetian Petterocer-atidae, which just reaches the lowest Frasnian, differs fromall other families in the initiation of additional lobes at theventer. The Devonopronoritidae (Bogoslovsky, 1958; 1969:290-295) contains solelyDevonopronorites, an endemic odd-ity from the Rudnyi Altai, Russia, with a bifid first laterallobe, an interesting approach to ceratitic frilling.

Here the name Pharciceratoidea is used in preference toTriainocerataceae essentially following A. K. Miller & W.M. Furnish (in Moore, 1957: L33), although the recogni-tion of the suborder Gephuroceratina rearranges the status.The superfamily Triainoceratoidea has been used by Bo-goslovsky (1969: 113) essentially as a replacement name forPharciceratoidea, which he had also used on earlier occasions(in Orlov, 1962: 341). The status of Triainocerae (Hyatt,1884: 333) is rather questionable for taxonomic purposes,and the form in which the name was introduced (withinbrackets rather than without as used for the clear designa-tions) implies that it was not envisaged as formal higher-taxon characterization (indeed, some groups proposed thuscomprise generic names only). It seems reasonable to preferthe unquestionably correct derivation of Pharciceratoidea.Notice, however, that the same comment could be made forTriainoceratidae, which is more recent, in its correct form,than Sandbergeroceratidae (Miller, 1938: 24, nom. transl.Ruzhencev, 1957: 55), so that there is a certain illogicalityin the procedure adopted here, the intention of which is todisturb terminology as little as possible.

In New York, only the late Givetian Pharciceratidae andthe Frasnian Triainoceratidae are known. The Eobelocerati-dae, Petteroceratidae, and Devonopronoritidae are notknown to be represented in North America.

Family PHARCICERATIDAE Hyatt, 1900Diagnosis.–Pharciceratoideans with 3 to at least 23 lateral

lobes (including those on the seam) in the adult with simpleto trifid ventral lobes. Growth lines biconvex. Form sub-globular to subevolute, outline usually rounded, rarely oxy-conic, mostly without ribs and nodes.

Discussion.–This family includes a wide range of distincttypes resulting from a radiation during the late Givetian fol-lowing initiation of forms with additional umbilical lobes,the characteristic of distinction from the supposed ancestralMaenioceratidae (House, 1978: 45).

The family includes the genera Pharciceras Hyatt, 1884(= Sphaeropharciceras Bogoslovsky, 1955b: 1104), Neophar-ciceras Bogoslovsky, 1955a: 95, StenopharcicerasMontesinos&Henn, 1986, and Synpharciceras Schindewolf, 1940: 430.It has been reviewed by Bogoslovsky (1969). Schindewolf(1940) distinguished a family, the Synpharciceratidae, andthis might prove useful if it is ever needed to separate mul-

tilobed forms, which could comprise these, from those withfewer lobes like Pharciceras. Too little is known, however, tofollow such a course at present and the Synpharciceratidaeis regarded as a junior synonym of Pharciceratidae.

Distribution.–Representatives known in Europe, NorthAfrica, Russia, China, and eastern North America (Becker &House, 2000).

Range.–In Europe, the family is restricted to late Givet-ian to earliest Frasnian: Pharciceras to Neopharciceras Geno-zones MD III-A-E to UD I-A. Formerly taken to berestricted to the long established lunulicosta Zone (lower-most Manticoceras Stufe), or Oberdevon I alpha, which is adivision now mostly in the Middle Devonian following thedecision of the Subcommission on Devonian Stratigraphyto raise the Middle/Upper Devonian boundary to a levelabove the range of the family (Klapper et al., 1987).

Genus PHARCICERAS Hyatt, 1884Type species.–Goniatites tridensG. & F. Sandberger, 1850,

by subsequent designation (Wedekind, 1918: 127).Diagnosis.–Pharciceratids with varied shell form, from

widely to narrowly umbilicate and well-rounded whorl sec-tion at least in early whorls. Growth lines biconvex with lap-pets often associated with ventrolateral furrows. Suture witha simple mid-ventral lobe, a first lateral lobe, in manysmaller than the second lateral lobe, and one to three lobesbetween the second lateral and the umbilical seam that formas umbilical lobes during ontogeny.

Included species.–Pharciceras tridens (Sandberger & Sand-berger, 1850), P. amplexum (Hall, 1886), P. bidentatum Pet-ter, 1959, and P. galeatum Wedekind, 1918.

Discussion.–With the creation of Stenopharciceras byMontesinos & Henn (1986), the genus is now restricted toforms with rather few lateral lobes. The simplest forms,Pharciceras tridens and P. amplexum, seem to be earliest; theformer is the earliest recorded in North Africa (Bensaïd,1974) and the latter, in New York, has the earliest conodontdate (late Middle varcus Zone).

Distribution.–Pharciceras is known from eastern NorthAmerica (Tully Fomation and equivalents between Senecaand Otsego counties, New York), Europe (Spain and Ger-many), and North Africa (see Addendum).

Range.–Givetian: Restricted to Pharciceras GenozoneMD III-A, which is defined by its appearance. Appears torange from top of conodont Middle varcus Zone to disparilisZone but range is uncertain, as conodonts are rare or absentat many of the localities.

Pharciceras amplexum (Hall, 1886)Pl. 1, Figs 1-8; Text-figs 26D-H; Table 2

Goniatites amplexus Hall, 1886, pl. 127 (12), fig. 1; Beecher inHall & Clarke, 1888: 39-40, pl. 127, fig. 1.

“Goniatites” amplexus. Cooper &Williams, 1935: 859.Maeneceras amplexum. Clarke & Ruedemann, 1903: 602.Manticoceras amplexum. Frech, 1913: 23.Manticoceras? amplexum. Miller, 1938: 74-76, pl. 13, figs 15-19.Pharciceras amplexum.House, 1962: 256, 272-274, text-figs 10C-

D, pl. 45, figs 1-4, 7-9; 1968: 1065; Bogoslovsky, 1969: 285;House & Ziegler, 1977: 90, text-fig. 91; House, 1979: 54,text-figs 9G-I; House & Kirchgasser, 1993: 276.

Type material.–The holotype (NYSM 3729), the onlyspecimen known to Hall when he illustrated the species, wasdescribed by Beecher (1888) and by Miller (1938) and isreillustrated here (Text-fig. 26F, Pl. 1, Fig. 3); it is a smallseptate specimen showing growth lines.

Additional material.–In addition to the material de-scribed by House (1962, USNM 143014, 137661, 96545aand b, 96552), material listed below now in the NYSM andSUI 37052a,b,c.

Dimensions.–See Table 2.Description.–Shell form subevolute; whorl outline in

early whorls depressed with well-rounded flanks, weak ven-trolateral spirals, and tabular venter. At approximately 25mm diameter, whorl height equals whorl width and approx-imately equidimensional outline continues to largest diam-eters seen (USNM 96545a), but WW/WH ratio varies. Inlarger specimens, ventrolateral groove better developed,often double (USNM 96545b; NYSM 11996 at 31 mm di-ameter) and well marked; less so in other cases (SUI37052c). Growth lines biconvex, passing slightly forwardover umbilical wall, then backward to broad, rounded sinuson mid-flanks, then sweeping forward into ventrolateral fur-row to form lappet and back into deep ventral sinus; lappetrelatively longer in earlier stages. Very fine ribbing developedon flanks before approximately 15 mm diameter: fine ribs

follow course of growth lines, seen on flanks only, number-ing 24 in previous half whorl at 13.6 mm diameter (SUI37052b) but coarser in other specimens (NYSM 12001).Suture as illustrated (Text-figs 26D-F); dorsal suture un-known.

Discussion.–Substantially more material is now availableon which to describe the species. The material is not wellpreserved and although commonly showing sutures andgrowth lines, detailed elucidation of ontogeny has not beenpossible. Many specimens are distorted as result of fractur-ing. Hence some of the measurements are approximate. Thisspecies differs from openly umbilicate maenioceratids onlyin the additional lobe near the umbilical seam. It is the ad-dition of one or more extra lobes in this position that is thegeneric distinction of Pharciceras from Maenioceras. Thespecies is very close to the lectotype, here designated, ofPharciceras tridens (G. & F. Sandberger, 1850: pl. 4, fig. 2;1851: 66; and refigured here, Pl. 1, Fig. 10). As can be seen,however, from the dimensions of the lectotype of P. tridenspreviously published (House, 1962: 273), that species ismore depressed at comparable diameters, although SUI37052a approaches it in certain respects. German materialof P. tridens (e. g., see Kullmann & Ziegler, 1970: text-fig.2A, pl. 1, fig. 3) shows that the type species can be consid-erably larger and in another specimen, illustrated by theSandberger brothers (1851: pl. 9, figs 2, 2a, 2b; refiguredby House & Ziegler, 1977: pl. 5, figs 24-26), the sutural el-ements seem more advanced at comparable diameters. Bothcharacters might suggest a later derivative and youngerspecies than P. amplexum.

The entry of Pharciceras was taken in Germany to markthe base of the Adorfian and Upper Devonian (Frech,1887). The attempt of Kullmann & Ziegler (1970), how-ever, to avoid assignment of P. amplexum to Pharciceras hasbeen precluded by the evidence from excellent material in

Table 2. Biometric data for Pharciceras amplexum (Hall, 1886) from the Tully Limestone. Here, and in all tables of biometric data tofollow, D = diameter, U = umbilicus, UW = umbilical width, Wh = internal whorl (aperture) height, WH = whorl height, WW = whorlwidth (Text-fig. 25). All measurements in millimeters.

Specimen D WW WH UW WW/WH UW/D

NYSM 3729 (holotype) ca. 17.5 8.5 6.0 10.0 1.42 0.57USNM 96545a 39.6 12.3 12.5 18.0 0.98 0.45USNM 96545b ca. 36.0 ca. 14.0 ca. 12.6 — 1.11 —USNM 96552c 33.4 ca. 11.5 ca. 11.0 14.3 1.05 0.43USNM 143014 ca. 30.5 11.0 10.0 ca. 13.3 1.10 0.44SUI 37052c ca. 29.0 ca. 11.7 — 12.8 — 0.44USNM 137661 29.0 9.7 10.0 12.3 0.97 0.42NYSM 11998 ca. 25.0 8.8 8.6 — 1.02 —NYSM 11997 22.6 8.6 — — — —SUI 37052b 16.0 ca. 7.0 ca. 5.0 — 1.40 —USNM 143014 ca. 15.0 7.7 4.8 6.4 1.60 0.43SUI 37052c 13.1 7.5 — — — —


the Iowa collection (House, 1978: 554). House (1962: 273)drew attention to the affinities of P. amplexum to Triainocerasbut the lack of marked ribbing precludes reference to theTriainoceratidae. Kullmann & Ziegler (1970) also consid-ered Tamarites (Bogoslovsky, 1969: 275), an evolute ribbedform that has a similar subdivision of the ventral lobe (butsimpler overall suture), and used evidence from G. A.Cooper’s comments on brachiopods associated withTamarites to argue that such types could be Middle Devon-ian – all without drawing attention to the fact that inCooper’s view of the placing of the Middle/Upper Devonianboundary (Cooper et al., 1942; Cooper, 1968), any I alphagoniatite would be assigned to the Middle Devonian becausehe took the Genundewa Limestone as the base of the UpperDevonian. All this has been resolved by the decision of theInternational Union of Geological Sciences (IUGS) andSubcommission on Devonian Stratigraphy (SDS) to definethe Middle/Upper Devonian boundary at a much higherlevel than the Tully Limestone (Klapper et al., 1987).

Distribution.–Moravia Bed or West Brook Member ofthe Tully Formation between Seneca and Cortland counties.Almost all of the known specimens of Pharciceras amplexumcertainly come from a single level in the Moravia Bed (Text-

figs 5-6) of the Tully Limestone (Heckel, 1973) or the upperpart of Bed G in the West Brook Member of Cooper &Williams (1935: 791). From Lodi Glen, Seneca Lake(Seneca County; Loc. 3), the most westerly known record,comes the holotype (NYSM 3729) and other specimens(NYSM 12002a and b, 12003) one of which was collectedin situ by J. W.Wells. From June’s Quarry, Tully (OnondagaCounty; Loc. 7), comes USNM 143014, SUI 37052a-c,and NYSM 11996-11999, 12004, 12005). From TinkersFalls (Onondaga County; Loc. 8), comes USNM 137661.

Also from the West Brook Member, but at localities far-ther east, are several specimens referred here that were col-lected during the survey by Cooper &Williams (1935). Theprecise correlation of these with the Pharciceras-bearing levelfarther west can only be inferred. From near Lebanon(Madison County; Loc. 9c), is a single specimen (USNM96552). From near Laurens (Otsego County; Loc. 9b), areseveral specimens (USNM 96545a and b). This locality nearLaurens is the most easternly record of P. amplexum known.There is no other record of this species elsewhere in easternNorth America of which we are aware. Molds of probablePharciceras occur in some abundance in the Filmore GlenBed of the Tully and rarely in the overlying Geneseo Shale

Text-fig. 26. Sutures and cross sections of Pharciceras tridens (G. & F. Sandberger, 1850) from Germany and the sutures and cross sectionof P. amplexum (Hall, 1886) from the Tully Limestone of New York. A-C. Pharciceras tridens. A. SMH 9935, suture of a specimen from theUntere Pharciceras-Lage of Kullmann & Ziegler (1970: 74) in the Martenberg section at Adorf (now Diemelsee), Germany, X 1.4. B-C.Sutures enlarged from the original figures of G. & F. Sandberger (1849-1856: pl. 4, fig. 4e, pl. 9, fig. 2b) from Germany; the sutures havebeen developed by abrasion that has resulted in the rounding of some of the lobes. D-H. Pharciceras amplexum. D. SUI 37052, suture of aspecimen from June's Quarry, Tully (Loc. 7) drawn by W. M. Furnish, X 3. E, USNM 143014, suture based on a specimen from the samelocality, X 3. F. NYSM 3729, suture based on the holotype from near Lodi Landing (Loc. 3), Seneca Lake, Seneca County, X 3. G, NYSM12002, reconstructed cross section based on a specimen from June's Quarry (Loc. 7), Tully, Onondaga County, X 1.8. H. NYSM 11999,reconstructed cross section based on a specimen from the same locality, X 1.8.



(Pl. 31, Figs 14-15).Range.–Givetian: Pharciceras Genozone MD III-A. Re-

gional Zone of P. amplexum (13). Conodont late Middle toUpper varcus Zone (Klapper, 1981: fig. 2). See also Abous-salam & Becker (2001: fig. 1) and Aboussalam (2003: fig.2).

Family TRIAINOCERATIDAE Hyatt, 1884Diagnosis.–Pharciceratoideans with four to perhaps ten

or so ventral lobes in adult with ventral lobe small and sim-ple to large and trifid. Shell form with strongly costate, evo-lute inner whorls, smoother outer whorls, and tendencytoward compressed or oxyconic form in adult.

Discussion.–The species included in the Triainoceratidaehave markedly ribbed inner whorls but costation is virtuallylost in adults. Almost all species show ventrolateral furrowsat some stage, and they are often double. There is a charac-teristic increase in the size of lobes from the umbilical seamtoward the venter. The critical distinction between the in-cluded genera, as presently understood, lies in the form ofthe ventral lobe. Much has to be learned concerning thefamily and, apart from Triainoceras gerassimovi Bogoslovsky,1958, none of the included species can be said to be wellknown.

The family Triainoceratidae (nom. correct., Ruzhentzev,1960: 175, pro Triainocerae Hyatt, 1884: 336) comprisesthe four genera Triainoceras, Sandbergeroceras, Wellsites, andSchindewolfoceras; all genera are confined to Oberdevon I ofthe traditional German definition. To these four genera istentatively added Tamarites Bogoslovsky, 1965, the stronglyribbed, serpenticonic form with few sutural elements, whichcould be the Middle Devonian ancestral representative.

Triainoceras is known from several species. The typespecies, by monotypy, is Goniatites costatus d’Archiac & deVerneuil (1842: 340, pl. 31, figs 1, 1a) from Eibach andOberscheld, Germany; the characters of the species wereworked out by Denckmann (1903b). T. gerassimovi Bo-goslovsky (1958: 127, pl. 8, fig. 3; 1969: 279, pl. 25, fig. 3),from the Rudnyi Altai, Russia, is known in considerable on-togenetic detail. Uncertainly placed here (because sutures inthem are not known) are T.? incertum (d’Archiac & deVerneuil, 1842: 342, pl. 2, figs 6, 6a) from Brilon, Germany,and T.? tuberculosum (d’Archiac & de Verneuil, 1842: 342,pl. 26, figs 4, 4a, 4b) from Oberscheld, Germany.

Miller (1938: 24, 178) grouped Sandbergeroceras, Schin-dewolfoceras, and Pseudarietites as a new subfamily, the Sand-bergeroceratinae. He considered Triainoceras as a synonymof Sandbergeroceras and the same view has been put to us byR. T. Becker; this is likely, or even probable. But it cannotbe accepted until detailed ontogenies have been describedof type material. It is now recognized that the Lower Car-

boniferous Pseudarietites is unrelated to these other generaand is derived from Gattendorfia in the early Tournaisian(Weyer, 1972). Miller (1938: 179) thought that Sandberge-roceras sandbergerorum Miller, 1938, from Oberscheld andKönigzug, Germany, shows evidence of an incipient trifidlobe, but now that the type has been photographically illus-trated for the first time (House & Ziegler, 1977, pl. 6, figs16-18) there is really no strong evidence for this. Certainlythere is nothing to approach the marked trifid lobes of T.gerassimovi and T. costatum (d'Archiac & de Verneuil, 0000),so that the generic distinction is worth keeping.

The material described here from New York is mostlypoorly preserved, very little of it showing evidence of su-tures. It appears, however, that shell ornament gives someconsistent stratigraphical information. None of the materialassigned to Sandbergeroceras shows a complete suture or ev-idence of the ventral lobe. Forms assigned to Sandbergero-ceras occur early, apparently in the Sonyea Group, probablybeginning in the Middlesex Shale. They are characterized byribbing in the inner whorls that is not known to extend be-yond 30-40 mm diameter, with individual ribs prorsiradiateto approximately rectiradiate (as in the type species of thegenus, S. sandbergerorum); fine growth-line lirae often lie be-tween the ribs. Specimens referred to Schindewolfoceras, onthe other hand, might begin in the upper Sonyea Group,but are characteristic of the West Falls Group. In the latter,the ribbing usually extends to at least 80 mm diameter, andthe course of the ribs is rursiradiate over the outer flanks andoften also distinctly backwardly directed over the lowerflanks. This is no more than an initial outline for discrimi-nation, but nothing more can be attempted with the mate-rial that is available.

Distribution.–Eastern North America, Europe, NorthAfrica, and Russia. It would seem that, on current evidence,Schindewolfoceras andWellsites are endemic to eastern NorthAmerica. The near restriction of the family to siltstone faciesin eastern North America is puzzling (Text-fig. 14).

Range.–European, North African, and Russian represen-tatives of the Triainoceratidae seem to be restricted to theFrasnian: Sandbergeroceras to BelocerasGenozones UD I-D-H.

Genus SANDBERGEROCERAS Hyatt, 1884Type species.–Sandbergeroceras sandbergerorum Miller,

1938, by subsequent designation (Miller, 1938: 178).Diagnosis.–Triainoceratids with evolute inner costate

whorls and ribbing, often to 40 mm diameter, with smooth,laterally compressed outer whorls often approaching dis-coidal form. Prominent growth-line lirae in inner whorls.Ribs prorsiradiate or approximately rectiradiate. Suture witha lobe centered on seam and four lateral lobes; ventral lobe

deep and flat-bottomed with small siphuncular lobe withinit; dorsal suture with mid-dorsal lobe and lobe between itand seam (full suture known only in the type species).

Included species.–Sandbergeroceras sandbergerorumMiller[1938: 178, proGoniatites tuberculoso-costatus (pars) G. & F.Sandberger, 1850]; S. syngonum Clarke (1897, nom. nud.;1898, 1899a-b: 106, pl. 7, fig. 19; NYSM 4077, lectotypehere chosen); S.? enfieldense n. sp. (see below).

Discussion.–Sandbergeroceras was erected by Hyatt (1884:333) with a misspelling as Sandbergeoceras (corrected byCrosse & Fisher, 1884: 415; ICZN, 1956; see Miller, 1952).From the originally mentioned included species, MillernamedGoniatites tuberculoso-costatus as type species, but be-cause this was an invalid combination of two of d’Archiac &de Verneuil’s specific names, Miller (1938) proposed S. sand-bergerorum as a replacement name for certain figures of G.& F. Sandberger (1850: pl. 4, fig. 1), which became the typespecies. Because it is not clear that the figure refers to onespecimen (although it might), House & Ziegler (1977: pl.6, figs 16-18) designated one specimen as lectotype and il-lustrated it photographically. The genus differs from Triain-oceras notably in the nature of the ventral lobe. Miller(1938), following Clarke (1897), but with a mark of doubt,referred S. syngonum from New York to the genus.

As applied to New York material, there is bound to bedoubt surrounding the use of this generic name because noNew York specimen referred here shows a complete suture.All assignments are therefore tentative.

Distribution.–Eastern North America (Sonyea Group,New York, and ?West Virginia) and Europe (Germany). InNew York, from the Sonyea Group (Middlesex and?Cashaqua Shale) in Ontario County and equivalent level(s)(Middlesex Shale and Cashaqua Shale) in Tioga County.The West Virginia record (Miller, 1938) remains uncon-firmed. In Germany, only the type material of the Sand-berger brothers exists from the Rhenish Schiefergebirge at alevel not documented.

Range.–Sandbergeroceras Genozone UD I-D to Probelo-ceras Genozone UD I-E, based on New York evidence.

Sandbergeroceras syngonum Clarke, 1897 (Group)Pl. 2, Figs 4-7, Pl. 3, Figs 5-6; Text-figs 27G-H; Table 3

Goniatites chemungense Clarke, 1885: 51.Sandbergeroceras syngonumClarke, 1897: 53; 1898, 1899a, b: 106-

108, ?text-figs 79-81, pl. 6, ?fig. 23, pl. 7, fig. ?19, fig. 20;House & Kirchgasser, 1993: 276.

Sandbergeroceras? syngonum. Miller, 1938: 179-182, pl. 36, figs 9-11.

Type material.–Clarke illustrated and mentioned onlyfour specimens in describing this species. A specimencrushed flat on a bedding surface (first mentioned in 1885according to Clarke, 1898: 108, footnote) is here selectedas lectotype (NYSM 4077, refigured here Pl. 3, Fig. 5).Some doubt must surround the determination of the othermaterial that is assigned to the species with a question mark.There is a protoconch and initial whorl (NYSM 4078) anda larger juvenile baritic replacement (NYSM 4075, refiguredhere, Pl. 2, Fig. 6) and a larger gutta percha mold referred toby Clarke as “probably” of the species (NYSM 4076, refig-ured here, Pl. 2, Figs 4-5). These comprise the type series.

Additional material.–Only three additional specimensare available, one collected by H. S.Williams from a localityinTiogo County, south of Harford Mills (Cortland County;Loc. 64), preserved as a partial external mold (NYSM12026, figured here Pl. 2, Fig. 7) and two loose specimens(NYSM 16577 and 16578) found by us at the same locality.

Dimensions.–See Table 3.Description.–Lectotype subevolute, but whorl section

cannot be determined. Ribs number approximately 14 inprevious half whorl at 43 mm diameter; ribs approximatelyrectiradiate but with slight flexure. One new specimen(NYSM 12026) shows 15 ribs in previous half whorl at ap-proximately 28 mm diameter. For earlier whorls, gutta per-cha mold (NYSM 4076, Pl. 2, Figs 4-5, Text-figs 27G-H)shows that early whorls have ribs prorsiradiate on flanks thatbecome almost rectiradiate by 19 mm diameter. Relation ofribs to growth-line lirae well shown by NYSM 4076 and byHarford Mills specimen (NYSM 12026); both showing liraeconforming with course of ribs and covering shell both overribs and between them with approximately 8-10 lirae from

Table 3. Biometric data for Sandbergeroceras syngonum Clarke, 1897, from the Middlesex Shale.


NYSM 4077 (lectotype) 52.0 — — — — —NYSM 12026 ca. 28.0 — 13.0 — — —NYSM 4076 19.0 — — ca. 9.8 — 0.52

ca. 15.0 6.3 4.6 — 1.37 —NYSM 4075 5.0 — — — — —

4.3 — 0.9 2.3 — 0.53


crest to crest.Nothing need be added to the original description of the

inner whorls given by Clarke (summarized by Miller, 1938)except to remark that their relationship to the larger speci-mens is inferred only. No adult sutures are known.

Discussion.–All of this material leaves much to be desired,but a distinctive pattern of ribs and growth-line pattern pro-vides a means of distinguishing the species. Sandbergerocerassandbergerorum differs in several respects from S. syngonum,notably in the absence of marked ventrolateral furrows and

Text-fig. 27. Sutures of Triainoceratidae and Beloceratidae of the late Devonian of various localities. A.Wellsites tynaniHouse & Kirchgasser,1993; SUI 42418, suture based on a specimen from the Rhinestreet Shale at Elmira (Loc. 70), Chemung County, X 1. B.Wellsites williamsi(Wells, 1956); NYSM/CU 40023, suture of the holotype, at whorl height ca. 72 mm, from Rhinestreet Shale equivalents at Bald Mountain(Loc. 68), Tioga County, X 1. C. Beloceras sagittarium (G. & F. Sandberger, 1851); suture of the type species of Beloceras based on a specimenfrom the Frasnian of Western Australia figured by Glenister (1958). D. Triainoceras gerassimovi Bogoslovsky, 1958; diagram illustrating theadult suture based on a specimen from the Rudny Altai, Russia, figured by Bogoslovsky (1969). E. Triainoceras costatum (d'Archiac & deVerneuil, 1842), the type species of the genus; suture reconstructed from the figure of the holotype from Oberscheld (Eibach), Germany. F.Sandbergeroceras sandbergerorumMiller, 1938; suture of the type species of the genus based on a specimen fromOberscheld (Königzug), Ger-many, figured by G. & F. Sandberger (1850). G-H. Sandbergeroceras syngonum Clarke, 1897; NYSM 4076, based on a specimen "from aconcretion on Honeoye Lake" and possibly from the Cashaqua Shale. G. Growth lines, X 9. H.Whorl cross section, X 2.5. I. Schindewolfoceraschemungense (Vanuxem, 1842); NYSM 4073, suture (reversed for comparison) based on the holotype, at ca. 73 mm diameter fromRhinestreet Shale equivalents near Owego (Loc. 69), Tioga County, X 2. J. Schindewolfoceras aff. chemungense; SUI 40080, suture of a spec-imen fromHwy 19 roadcut near Big Spring, Missouri, collected by J. Ragan; from float on a Callaway Limestone exposure but it seems prob-able that it was derived from the overlying Burlington Limestone, X 3.5.



sharp ventrolateral shoulders at comparable diameters (cf.,House & Ziegler, 1977: pl. 6, fig. 16) and the narrower ven-tral sinus band; ribs in the German specimen number ap-proximately 12 in the previous half whorl at 28 mmdiameter and they seem more sharply defined than in S. syn-gonum.

Distribution.–Middlesex Shale (Sonyea Group) in On-tario or Yates county and Tioga County. Possibly in theCashaqua Shale (Sonyea Group) in Ontario and Yates coun-ties. The record from Tygart Valley (Randolph County),West Virginia, is unconfirmed (Miller, 1938; House, 1978).

The lectotype (NYSM 4077) is recorded as from the“lower black band” from Snyder’s Gully, Naples, a creek thatcrosses the Ontario-Yates county line near Woodville (On-tario County; Loc. 46c); the preservation and lithology con-firm that it came from the Middlesex Shale. The followingspecimens reported from the Portage or Naples beds couldhave come from the Cashaqua Shale: NYSM 4075, a bariticspecimen from a concretion at Honeoye Lake (OntarioCounty); NYSM 4076, a gutta percha mold of a specimenfrom a crinoidal concretion at Naples (Ontario County);NYSM 4078, a baritic specimen from possibly the samecrinoidal horizon at Naples. Although the labels withNYSM 4076 and 4078 refer to their source as “crinoidalblocks,” which could be from the Cashaqua, there is somesuggestion of High Point Sandstone lithology (Clarke &Luther, 1904). Naples is the type locality for the High PointSandstone, a unit higher up in the West Falls Group.

The Harford Mills (Loc. 64) specimen (NYSM 12026)fromTioga County collected by H. S. Williams is probablyfrom an upper Middlesex Shale equivalent. In 1998, wefound two loose fragments of the species (NYSM 16577 and16578) in siltstone and silty mudrock within a section ofblack shale at the locality.

Because this is the only distinctive form known from theMiddlesex Shale, the Regional Zone of Sandbergeroceras syn-gonum refers to this level and up to the entry of the distinc-tive Cashaqua Shale faunas with the next zone fossil,Probeloceras lutheri. It could be that the range of S. syngonumcontinues into the next zone or higher, but that is uncon-firmed.

Range.– Frasnian: Sandbergeroceras Genozone UD I-D.Regional Zone of Sandbergeroceras syngonum (18).

Sandbergeroceras? enfieldense n. sp.Pl. 2, Figs 1-2, Pl. 3, Figs 2-3

Sandbergeroceras? enfieldense Kirchgasser &House, 1981: 48, nom.nud.; House & Kirchgasser, 1993: 276.Etymology.–Named for the Enfield Formation, near

Ithaca (Tompkins County), the source of the holotype

(NYSM 12028).Material.–Two large specimens recorded as from the

“Enfield” Formation: the holotype (NYSM 12028) from theHungerford (University) Quarry (Loc. 66), near Ithaca, andanother specimen (a plastotype) (NYSM 12027; PRI 8742)from Bald Hill, Caroline (Tompkins County; Loc. 94). Bothare preserved mostly as external molds in fine siltstone.

Diagnosis.–Species of Sandbergeroceras in which ribbingis lost by approximately 20 mm diameter and ribs becomeless frequent just before their loss. Adult much larger than inany other known member of genus.

Dimensions.–NYSM 12028 (holotype), D = ca. 150.0mm, WH = ca. 68.0 mm, Wh = ca. 46.0 mm, UW = 48.0mm, UW/D = 0.32; NYSM 12027 (plastotype), D = ca.140.0 mm, UW = 39.0 mm, UW/D = 0.28.

Description.–Inner whorls strongly costate with ribsnumbering 12 in previous whorl at umbilical width of 17mm in holotype and 11 at same umbilical width in plasto-type. Ribs rectiradiate where best developed but in plasto-type, as ribs are lost they are progressively more prorsiradiateand follow course of growth lines that sweep forward fromseam but are more rectiradiate past the not-very-well-marked umbilical shoulder. Adult whorls smooth but withslight nodes replacing ribs after approximately 20 mm di-ameter. Adult laterally compressed, perhaps suboxyconic.

The holotype is a phragmocone; the diameter of thecomplete conch could have exceeded 250 mm. It shows partof a suture at the near-maximum diameter, suggesting thepresence of approximately five or more lobes on the flanks.The saddle is well rounded and the lobe deep and almostparallel-sided with a pinched acute lobe.

Discussion.–These giant specimens differ from all knownmembers of the genus in their size and ribbing pattern. Theyshare giant size with Wellsites known from rather farthersouth, but that genus is oxyconic and has a closed umbilicusas nearly as can be determined. Even though the ventral su-ture is not known, the species is assigned, with question, toSandbergeroceras.

Distribution.–Known only from the Enfield Formationnear Ithaca (Tompkins County); horizon undetermined butprobably from equivalents of the upper Cashaqua Shale(Sonyea Group) and lower Rhinestreet Shale (West FallsGroup). The holotype is from the Hungerford (University)Quarry (Loc. 66), southeast of Ithaca, and the plastotype isfrom Bald Hill, Caroline (Loc. 94), southeast of Brookton-dale (Tompkins County). The innermost whorls describedunder Sandbergeroceras? syngonum might belong to thisspecies.

Range.–Frasnian: ?Probeloceras or Prochorites or Mesobe-loceras Genozone(s) UD I-?E, or F, or G. ?Regional Zone ofProbeloceras lutheri (19) or Prochorites alveolatus (20) or

Naplesites iynx (21).

GenusWELLSITES House & Kirchgasser, 1993Type species.–Wellsites tynaniHouse & Kirchgasser, 1993,

by original designation.Diagnosis.–Oxyconic triainoceratid without ribbing, ex-

cept perhaps in early whorls, with involute conch. Primaryventral lobe with single division and seven lateral lobes (in-ferred to have developed as umbilical lobes) with all ele-ments rather deep.

Distribution.–Lower Rhinestreet Shale (West FallsGroup; Tompkins and Chemung counties).

Range.–Frasnian: ?Mesobeloceras or Beloceras GenozoneUD I-?G, H. ?Regional Zone of Naplesites iynx (21a) orWellsites tynani (21b).

Wellsites tynani House & Kirchgasser, 1993Pl. 1, Fig. 9; Text-fig. 27A

Wellsites tynani House & Kirchgasser 1993: 273, fig. 5A.

Type material.–Holotype, SUI 42418.Diagnosis.–As for genus but with larger lateral lobes

swollen and terminations pinched.Description.–Holotype a large crushed specimen showing

phragmocone and inferred oxyconic form at 190 mm diam-eter. With body chamber, it would have exceeded 250 mmin diameter. Ventral suture with in excess of 10 lateral lobesof which second lateral is deepest; lateral lobes (U in origin)decrease in size toward umbilicus. Larger lobes showingpinched terminations. Inner whorls not visible. Suture as il-lustrated in Text-fig. 27A.

Discussion.–This form clearly belongs to the groupnamed Beloceras by Wells (1956b) but does not show thesigmoidal lobes and saddles of that genus; the swollen lobesand pinched terminations show that this specimen should bereferred to the Pharciceratoidea. It is presumed to be a tri-ainoceratid that has lost ribbing, either completely or shownonly in the innermost whorls, which are not seen in theholotype.

Distribution.–Lower Rhinestreet Shale (West FallsGroup; Chemung County). The holotype, collected by M.C.Tynan, is recorded from spoil in a roadcut for US Rte. 17,0.16 km east of the crossing of US Rte. 17 and WatercureRoad, Elmira (Chemung County; Loc. 70), at an altitudeof approximately 910 ft (277 m). The horizon is said to bethe Moreland Shale, the basal tongue of the RhinestreetShale, but see comments in locality description (Loc. 70).

Range.–Frasnian: ?Mesobeloceras or Beloceras GenozoneUD I-G or H. ?Regional Zone of Naplesites iynx (21a) orWellsites tynani (21b).

Wellsites williamsi (Wells, 1956)Text-fig. 27B

Beloceras williamsi Wells, 1956b: 751, text-figs 2A, C, pl. 82, fig.4; House 1962: 258.

Wellsites williamsi. House & Kirchgasser, 1993: 273, fig. 5B.

Type material.–A single incomplete specimen preservedas a partial internal mold is the holotype (NYSM/CU40023); only a small portion of the body chamber is pre-served.

Diagnosis.–Differing from only other species in genus,Wellsites tynani, in having longer sutural elements and ab-sence in them of swollen form and pinched terminations.

Description.–Holotype illustrated and described byWells(1956b), so only brief comment is required. Largest whorlheight seen is approximately 80 mm, with last septum beingat approximately 74 mm diameter. Only portion of umbili-cus preserved, but clearly open although small. Whorl sec-tions seem clearly oxyconic. Sutures (Text-fig. 27B) show atleast nine lateral lobes with second lateral deepest; sutureseems somewhat distorted but lobes subacute and saddlesrounded.

Discussion.–With the removal of this species from Belo-ceras, there is now no evidence for the Beloceras in New Yorkor North America.

Distribution.–Bald Mountain (Loc. 68) southeast ofIthaca (Tompkins County). The level, recorded as approxi-mately 150 ft (46 m) above the base of the Cayuta Shale,suggests a horizon above the Moreland Shale, but in any casean early Rhinestreet Shale (West Falls Group) equivalent.

Range.–Frasnian: ?Beloceras Genozone UD I-?H. ?Re-gional Zone of Wellsites tynani (?21b).

Genus SCHINDEWOLFOCERASMiller, 1938Type species.–Goniatites chemungensisVanuxem, 1842, by

original designation.Diagnosis.–Triainoceratids with evolute to subevolute

shells and ribbing normally to at least 80 mm diameter, withribs usually rursiradiate in outer whorls and with ventrolat-eral furrows. Suture with short, squat, V-shaped ventral lobeand approximately six lateral lobes decreasing in size towardumbilical seam. Dorsal suture unknown.

Included species:–Schindewolfoceras chemungense (Vanux-em, 1842: 182, text-fig. 49, 1); S.? equicostatumHall (1874:3-4; 1876: pl. 69, fig. 10); S.? aff. equicostatum (Hall, 1874).

Discussion:–Schindewolfoceras was proposed by Miller(1938: 181) with Goniatites chemungensis as the designatedtype species. He included in it the two named Chemungspecimens of Vanuxem and Hall that were, at that time, re-ferred to Prolecanites. The two type specimens are not wellpreserved; only the holotype of S. chemungense shows a su-


ture and this was clearly a large goniatite (holotype, NYSM4073, figured herein, Pl. 2, Fig. 8, Text-fig. 27I). Both typespecimens were described by Miller (1938). The genus andspecies included in it are unknown outside New York State.

No additional material has been found in this studyapart from the specimen referred to Schindewolfoceras? aff.equicostatum from Fairfield Forest (Loc. 67) near Speedsville(Tioga County; NYSM 12027, Pl. 3, Fig. 1), and anotherbut very poorly preserved specimen (NYSM 12025) fromnear Chemung (Chemung County; Loc. 95), figured here as?Schindewolfoceras sp. (Pl. 2, Fig. 3).

Distribution.–West Falls Group in Tioga and Chemungcounties. Both type specimens are from the “Chemung”Group and the localities suggest West Falls Group equiva-lents. Schindewolfoceras? aff. equicostatum at Loc. 67 (TiogaCounty) might be a mid-Rhinestreet occurrence.

Range.–Frasnian: ?Mesobeloceras to Beloceras GenozoneUD I-?G to ?H.

Schindewolfoceras chemungense (Vanuxem, 1842)Pl. 2, Fig. 8; Text-fig. 27I

Goniatites chemungensis Vanuxem, 1842: 182, text-fig. 49, fig. 1;Hall, 1876: pl. 69, fig. 9, pl. 74, fig. 6; 1879: 467-469, pl. 69,fig. 9, pl. 74, fig. 6; Hyatt, 1884: 333.

Schindewolfoceras chemungense. Miller, 1938: 182-184, fig. 38B,pl. 36, figs 7-8; Bogoslovsky, 1969: 282-283; House & Kirch-gasser, 1993: 276, 278.

Type material.– The holoype (NYSM 4073), figured byVanuxem & Hall, is poorly preserved in fine silstone; it islaterally compressed and distorted and the body chamber isnot preserved.

Dimensions.–The holotype (NYSM 4073): D = ca. 78.0mm, Wh = ca. 28.0 mm, UW = ca. 31.0 mm, UW/D =0.40; S. aff. chemungense (SUI 40080, Pl. 4, Figs. 1-4): D =41.2 mm, WW = 14.8 mm, WH = 15.4 mm, Wh = 10.3mm, UW = 16.1 mm, WW/WH = 0.96, UW/D = 0.39.

Description.–Holotype indicates rather evolute innerwhorls, becoming subevolute in outer whorls. Flanks appearto have been rounded and without marked umbilical shoul-der; venter likely well-rounded but might have approachedsubacute. No evidence of ventrolateral furrows but perhapsa feature of preservation. Ribs on inner whorls, at approxi-mately 22 mm diameter, 1.4 mm apart on mid-flanks, pro-gressively becoming less prominent in last whorl seen andnearly rectiradiate on low flanks; earlier with backward de-flection and in last whorl with backward deflection on outerflanks. Ribs becoming weaker toward venter and numberingapproximately nine in previous half whorl at ca. 70 mm di-ameter. Ventral sutures (Text-fig. 27I) comprising six laterallobes increasing regularly in size toward venter; outermost

becoming nearly parallel-sided and subacute; ventral lobebroadly V-shaped, making first lateral saddle asymmetric.

Discussion.–No new material has come to light nor hasanything comparable to this species been noted in otherareas. For present purposes, a difference in rib frequency isused to separate this species from Schindewolfoceras?equicostatum that has significantly more ribs at a comparablediameter. Because the specimen is still septate, large adultsize, in excess of 200 mm diameter, is indicated.

Distribution.–Rhinestreet Shale equivalent (West FallsGroup; Tioga County). Vanuxem did not give a precise lo-cality for his specimen, but Hall (1879: pl. 69, fig. 9) refig-ured it and gave the locality as “Chemung near Owego,N.Y.” This is our Loc. 69 (Tioga County). The horizon isthought to be an equivalent of the Rhinestreet Shale. Miller(1938: 184) gave the locality as “Cayuta Shale member(Spirifer disjunctus zone) of the Chemung Formation.”

Range.–Frasnian: ?Beloceras Genozone UD I-?H. Re-gional Zone of Schindewolfoceras chemungense (21c).

Schindewolfoceras? equicostatum (Hall, 1874)Pl. 3, Fig. 4

Goniatites Chemungensis var. equicostatus Hall, 1874: 3-4; 1876,pl. 69, fig. 10.

Sandbergeroceras Chemungense Clarke, 1898, 1899a, b (pars): 108.Schindewolfoceras? equicostatum.Miller, 1938: 184, pl. 36, fig. 12;

Bogoslovsky, 1969: 283; House & Kirchgasser, 1993: 276.

Type material.–Only known by the holotype (NYSM4074) preserved in buff-weathering siltstone as a crushed in-ternal and part external mold; incomplete and not showingsuture.

Description.–Adult subevolute; whorls with well-roundedconvex flanks passing to rounded, weakly developed umbil-ical shoulder. Evidence of double ventrolateral furrow, butventer not seen; maximum diameter seen approximately 70mm. Ribs well developed, forming and slightly prorsiradiateover umbilical shoulder, then passing slightly backward overflanks but lost after evidence of sharp forward projectioninto ventrolateral furrow. Ribbing almost completely lost inoutermost quarter whorl (seen as external mold). Ribs num-bering 14 in previous half whorl at 70 mm diameter.

Discussion.–This form differs from Schindewolfoceraschemungense in the greater frequency of ribbing and the ev-idence of ventrolateral furrows. In the absence of evidence ofsutures, the generic assignment must remain in doubt.

Distribution.–Unknown, but probably low in the WestFalls Group of southern Tioga or Chemung counties. Orig-inally said to come from Athens (Bradford County), Penn-sylvania, and later said to come from a boulder, and hencepresumably loose. This could explain subsequent references



to it as from New York State because Athens is only a fewmiles south of the state line near Waverley (Tioga County),and it might have been inferred to have been derived fromthe north. Hall (1879) considered that the brachiopods inthe block indicatd that it came from the lower beds of theChemung Group which are well developed in the vinicity ofWaverly and Athens. It seems likely that the level is the earlyWest Falls Group of modern terminology.

Range.–Frasnian: ?Mesobeloceras or Beloceras GenozoneUD I-?G or H. Regional Zone uncertain (21).

Schindewolfoceras? aff. equicostatum (Hall, 1874)Pl. 3, Fig. 1

Schindewolfoceras? aff. equicostatum. House & Kirchgasser, 1993:276.

Material.–A single specimen (NYSM 12027), preservedin fine shelly siltstone, without any trace of sutures and in-ferred to be a body chamber.

Description.–Maximum diameter (estimated) approxi-mately 45 mm with convex flanks, prominent ventrolateralgroove with slight raised ridges on either side, and sugges-tions of tabular venter. Ribs numbering approximately 11in previous half whorl at ca. 45 mm diameter; these fadingboth in outermost flanks and toward aperture. Ribs withslight backward deflection in earliest whorl seen but becom-ing rather more rectiradiate; with evidence of forward sweepof ribs, as they are lost, toward ventrolateral groove.

Remarks.–This is a very similar conch to Schindewolfo-ceras? equicostatum but differs in the rather earlier loss of rib-bing and in their lesser frequency. The latter is acharacteristic of S. chemungense, but the ventrolateral furrowis not (yet) known in that species.

Distribution.–Middle Rhinestreet Shale equivalent (WestFalls Group; Tioga County). The single specimen collectedby J. W. Wells is from Fairfield Forest (Loc. 67) nearSpeedsville (Tioga County), from the Upper Cayuta Shale atapproximately 400 ft (122 m) above a Tropidoleptus horizon.From a level between the Moreland Shale and Roricks GlenShale and thus a mid-Rhinestreet Shale equivalent.

Range.–Frasnian: Uncertain Genozone UD I-?G, H. Re-gional Zone uncertain (?21-22).

Superfamily GEPHUROCERATOIDEA Frech, 1897This major group, typical of the Frasnian and extinct by itsclose, is thought to have arisen from simple serpenticonicanarcestids. It includes the Acanthoclymeniidae, Koeneniti-dae, Gephuroceratidae, and Beloceratidae, all of which arewell represented in the New York faunas. The superfamilybecame extinct in the latest Frasnian Kellwasser Event.

Family ACANTHOCLYMENIIDAE Schindewolf, 1955This family is thought to have arisen from the Anarcestidae,particularly from the Atlantoceras group, by the division ofthe ventral lobe and the addition of other lobes between theprimary subumbilical lobe and the dorsal lobe. As presentlyconceived, the family comprises the genera AcanthoclymeniaHyatt, 1900, Pseudoprobeloceras Bensaïd, 1974, PonticerasMatern, 1929, Gogoceras Becker et al., 1993, Uchtites Bo-goslovsky, 1969, Prochorites Clausen, 1969 (= Eidoprobelo-ceras Kirchgasser, 1968, nom. nud.), and probably EnseitesBecker & House, 1993, and a recent new genus, ChutocerasBecker et al., 2000, based on material from the Timan-Pe-chora area of Russia. Acanthoclymenia, Ponticeras, Chutoceras,and Prochorites occur in North America. The family becameextinct with the Upper Kellwasser Event.

Distribution.–Eastern North America, Europe, Russia,North Africa, and Western Australia.

Range.–Upper Givetian to top of Frasnian: MD III-Dto UD I-L.

Genus PONTICERASMatern, 1929Type species.–Ammonites aequabile Beyrich (1837: 34) by

original designation.Diagnosis.–Conch evolute in early stages, subinvolute in

adult, usually laterally compressed, with phragmocone com-monly exceeding 100 mm diameter. Whorl section roundedto compressed. Venter tabular or rounded or oxyconic.Withor without ventrolateral furrows. Suture as in Acanthocly-menia but some with divided ventral saddle very large. Lat-eral saddle broadly arched, symmetrical to stronglyasymmetric with steep dorsal face; dorsal lobe deep. Laterallobe broadly to narrowly rounded, sometimes pointed.Growth lines strongly to weakly biconvex; in some,markedly rursiradiate across flanks with strong ventrolateralsalient.

Included species.–Ponticeras aequabile (Beyrich, 1837), P.barroisi (Wedekind, 1917, 1918), P. perlatum (Hall, 1874,1876), P. auritum (Holzapfel, 1899), P. bisulcatum (Keyser-ling, 1844), P. discoidale Glenister, 1958, P. domanicense(Holzapfel, 1899), P. kayseri (Wedekind, 1917, 1918), andP. keyserlingi (Holzapfel, 1899).

Discussion.–A neotype for Ponticeras aequabile waserected by House & Ziegler (1977: 77) on the assumptionthat Beyrich’s specimen had been lost, it not having beennoted for a century and a half. The neotype was designatedas a specimen that had been figured byWedekind (1918: pl.21, fig. 6) as the species (it was refigured by House &Ziegler, 1977: pl. 1, fig. 25). R. T. Becker recently discoveredBeyrich’s specimen in the Museum für Naturkunde, Berlin.The designation of the neotype therefore is invalidated. R.T. Becker is preparing a description of the specimen that one

of us (MRH) has examined. Beyrich’s original descriptionand illustration are good and the specimen shows a whorlsection with a narrowly tabular venter. In this it differs frommost New York specimens but agrees with the many, usuallylarger, specimens that Bogoslovsky (1969) described fromthe Timan-Pechora region of northern European Russia. Itraises the question as to whether earlier members, such as P.perlatum, with rounded venters, should in the future be tax-onomically distinguished from a later group with tabularventers.

Distribution.–Late Middle Devonian to mid-Frasnian.MD III-D to UDI-E.

Ponticeras perlatum (Hall, 1874)Pl. 4, Figs 5-9, Pl. 5, Figs 1-9;

Text-figs 28A-G, 29A-C, 32A; Table 4

Goniatites complanatus perlatus Hall, 1874: 1-2; 1875: 132-134;1876: pl. 70, fig. 12; 1879: 458-459, pl. 70, fig. 12.

Goniatites complanatus (Hall, 1874). Hall, 1879 (in part): 455-457, pl. 70, figs 6-8.

Manticoceras perlatum. Miller, 1938: 95-97, pl. 24, figs 11-12.Ponticeras perlatum.Wells, 1956a: 748; House, 1962: 256, pl. 45,

figs 10-12, text-figs 3A-B; Kirchgasser, 1975: 86-88, pl. 3, figs6-11, text-fig. 11, table 4; House & Kirchgasser, 1993: 274-276.

Type specimen.–NYSM 3649, specimen figured by Hall(1876: pl. 70, fig. 12; 1879: pl. 70, fig. 12) and herein (Pl.4, Fig. 9) from the Sherburne Siltstone, Genesee Group,Homer (Cortland County).

New material.–New specimens from the Lodi Limestoneat Romulus Town Quarry (Loc. 27a), Ovid, and one fromthe Lodi at Beards Creek (Loc. 18/4) were added to the col-lections in the NYSM and the CUmade by various workersover the past century from the Geneseo Shale, Lodi Lime-stone and Sherburne Siltstone from localities betweenSeneca and Cayuga lakes and eastward to Homer, Cortland,

andTruxton (Cortland County).We also include a few spec-imens from more eastward localities collected by CharlesThayer during a study of the paleoecology of the GeneseeGroup (Thayer, 1974). Most of the Thayer material ispoorly preserved molds but there is a fine specimen from atongue of black shale (Geneseo Shale) from near Upperville(Chenango County; Thayer’s loc. A-9051) and two speci-mens from the Sherburne Siltstone from localties southeastof Truxton (Cortland County; Locs A9157 and A9158).Thayer’s collection will be deposited at the NYSM.

Dimensions.–See Table 4.Diagnosis.–Adult shell large, to 90 mm diameter, evolute

and compressed, with flanks curving gradually to weak ven-trolateral furrows and narrowly rounded venter. Adult suturewith broadly rounded median saddle, deeply pointed ven-trolateral lobe, high, broadly rounded and slightly asymmet-ric lateral saddle, and broadly rounded lateral lobe. Growthlines strongly sinuous in adult with prominent, roundedsalient, dorsad of ventrolateral furrow, which appear as lowarcuate ribs on internal molds (Pl. 5, Figs 8-9). Growth-linelirae on flanks of inner whorls relatively straight and fascic-ulate, becoming more sinuose and evenly spaced. See alsoKirchgasser (1975: 86-88).

Discussion.–Ponticeras perlatum enters the New York sec-tion near the top of the Geneseo Shale in the form of mostlycrushed specimens at localities near Cayuga Lake (TompkinsCounty; Pl. 4, Fig. 6). Better specimens with preserved shellsare in the main bed of the Lodi Limestone near the base ofthe overlying Sherburne Siltstone in the Seneca and Cayugalake valleys, and especially from Lodi Glen (Loc. 27) nearLodi (Seneca County), and Romulus Town Quarry (Loc.27a), near Ovid (Seneca County). Fragments of P. perlatumhave been noted in the nodular westernmost bed of the Lodiat Beards Creek (Loc. 18/4) in the Genesee Valley (Text-fig.28F). A specimen with a rather narrow lateral lobe from thethird cycle of the Lodi Limestone higher in Sherburne atSeneca Lake (Loc. M, Ov-13) is referred to P. cf. perlatum

Table 4. Biometric data for Ponticeras perlatum (Hall, 1874) from the Lodi Limestone. See Text-fig. 32 for graphs.


NYSM 12166 ca. 92 ca. 15 ca. 34 ca. 32 0.44 0.35ca. 63 ca. 11 ca. 25 ca. 22 0.44 0.35ca. 45 ca. 9 ca. 16 ca. 16 0.56 0.36

NYSM 13173 60.8 11.3 24.3 19.3 0.47 0.3241.5 9.1 17.2 12.4 0.53 0.3028 ca. 6.0 11.9 7.7 0.50 0.288.2 2.5 3.5 2.8 0.71 0.345.6 1.9 2.0 2.2 0.95 0.394.3 1.4 1.4 1.7 1.00 0.403.4 ca. 1.1 1.1 1.3 1.00 0.382.5 ca. 0.90 0.83 0.90 1.08 0.36


Text-fig. 28. Sutures and cross sections of Ponticeras and Chutoceras from the Genesee Group in New York. A-G. Ponticeras perlatum (Hall,1874). A. NYSM 5122, suture and umbilicus, reversed, based on a specimen figured by Clarke (1898: pl. 24, fig. 12) from the SherburneSiltstone at Ithaca, Tompkins County (from House, 1962: text-fig. 3B), X 1.1. B, G. NYSM 13173, suture and cross section at 61.3 mmdiameter, based on a specimen from the Lodi Limestone at Ovid (Loc. 27a, WTK Loc. 3012), Seneca County (from Kirchgasser, 1975), X1.7. C-D. NYSM 13172, suture and cross section at ca. 41 mm diameter, based on a specimen from the Lodi Limestone at Lodi Glen (Loc.27/1; WTK Loc. 3013), Lodi Glen, Seneca County, X 1.7. E. NYSM/CU 41053, suture and growth line (dotted) of a specimen from theLodi Limestone, Hubbard Quarry (Loc. 27b), southwest of Kidders, Seneca County. F. NYSM 16567 [3852/1], cross section based on aspecimen from the Penn Yan Shale (Loc. 18/4), Beards Creek, Leicester, Livingston County. H-K. Chutoceras nundaium (Hall, 1874). H.NYSM 3754, a specimen figured by Hall (1888: pl. 128, fig. 2) from the Ithaca Formation, University Quarry (Loc X, Dy-10), Ithaca, Tomp-kins County, X 1.7. I-J, NYSM 3753, suture and cross section of a specimen figured by Hall (1888: pl. 128, fig. 1) from the same locality,X 1.1. K. NYSM 3751, suture of the lectotype (here designated) figured by Hall (1876: pl. 70, fig. 15) from the Ithaca Formation, SouthHill (Loc. AA, Dy-10c), Ithaca, Tompkins County, X 0.8. Scales = 5 mm (F); 1 cm (E, H-I).


Text-fig. 29. Sutures and cross sections of Ponticeras spp. from the Lodi Limestone (lower Penn Yan Shale) of New York State. A-C. Ponticerasperlatum (Hall, 1874). A. NYSM 12166, cross section of a specimen from the Lodi Limestone (Lodi B) at RomulusTownQuarry (Loc. 27a/1;WTK 3012), Ovid, Seneca County, X 1.8. B-C. NYSM 12167, cross section at ca. 27 mm whorl height, and suture (reversed) at ca. 24mm whorl height, based on a specimen from the Lodi Limestone (Lodi B) from the same locality, X 3.6. D-E. Ponticeras cf. perlatum, NYSM12168, cross section and suture (reversed) at ca. 30 mm whorl height, of a specimen collected by G. Baird from the third cycle of the LodiLimestone (Lodi C), 5.5 m (18 ft) above the main Lodi Limestone (Lodi B) at Loc. M (Ov-13), a gully 0.3 mi (0.48 km) north of TommyCreek, 2.6 mi (4.2 km) northwest of Lodi, Seneca County; loc. 18 of Baird et al. (1989), X 1.8. Scales = 1 cm (A), 5 mm (B-E).



(Text-figs 29D-E).Ponticeras perlatum was collected or noted from a num-

ber of localities and horizons within the Sherburne Siltstonein the Cayuga Valley by several workers beginning in thenineteenth century, most notably Williams (1884), Kindle(1896), and Boekenkamp (1963) (see locality discussions).Most specimens are poorly preserved and correlations withspecific horizons within the Lodi and Penn Yan to the westhave not been established (see Baird et al., 1989). Similarlythe records of P. perlatum from Cortland County are poorlyconstrained stratigraphically, but all are presumably fromthe upper Geneseo Shale or Sherbrune Siltstone or equiva-lents.

Distribution.–Uppermost Geneseo and lower Penn Yanshales (and equivalent Sherburne Siltstone to base of Ren-wick Shale) (Genesee Group), from Livingston to Cortlandcounties. Although Ponticeras can occur lower down in theGeneseo Shale (Geneseo Limestone Horizon and Fir TreePyrite), undoubted P. perlatum first occurs in the uppermostGeneseo Shale (Givetian conodont disparilis Zone) andranges upward into the Lodi Limestone (Givetian conodontnorrisi Zone) in the lower Penn Yan Shale and equivalentSherburne Siltstone. In the Cayuga Valley, P. perlatum rangesto the top of the Sherburne Siltstone at its contact with theoverlying black Renwick Shale, beds that correlate westwardto horizons with lower Frasnian MN Zone 1 conodonts. AtAbbey Gulf (Loc. 21), in the Honeoye Lake Valley, Pon-ticeras occurs in Penn Yan Black Shale B (above the Lodi andbelow the Renwick), a level in MN Zone 1 (Kirchgasser,1994: fig. 4). Farther west, in the Genesee Valley, P. perlatumcan range higher in the Penn Yan to levels above the SBblack shale horizon (conodont MN 1 Zone) but probablydoes not overlap the lower range of Koenenites (conodontMN Zone 2).

Range.–Uppermost Givetian to lower Frasnian: lowerPonticeras or PetterocerasGenozone MD III-E to upper Pon-ticeras or Neopharciceras Genozone UD I-A. Regional Zoneof P. perlatum (15b). Conodont disparilis and norrisi Zonesof Givetian to lower falsiovalis and MN Zone 1 of Frasnian(Kirchgasser, 1994).

Ponticeras? sp.Text-fig. 33M

Archoceras (Atlantoceras) sp. Kirchgasser & House, 1981: 42-43.Manticoceras cf. evolutum Petter, 1959. House & Kirchgasser,

1993: 276-277.

Material.–NYSM 12169, a calcitic replacement from theBeards Creek Horizon (18a/5), and six calcitic replacementsfrom the Fossil Log Horizon (18a/4b) (Williamsburgh Bedof Over et al., 1999, 2003) in the West River Shale, 2.4 m

and 3.4 m below the base of the Middlesex Shale, at BeardsCreek (Loc. 18), near Leicester (Livingston County).

Discussion.–The shells are juvenile stages and are toosmall to describe accurately. The preserved inner whorls areevolute to serpenticonic and distinctly more rounded anddepressed than those of specimens of Acanthoclymenia aff.neapolitana from the Fossil Log Horizon, 3.4 m below thebase of the Middlesex. Previously this form was referred toArchoceras (Atlantoceras) sp. (Kirchgasser & House, 1981).In Archoceras, however, lateral furrows are typically present.There is some indication that the mid-ventral lobe is divid-ing. Consequently we designate the specimens as Ponticeras?sp. It is now considered that this material cannot be referredto Manticoceras evolutum (see House & Kirchgasser, 1993:276-277).

Distribution.–Uppermost West River Shale (GeneseeGroup; Livingston County).

Range.–Frasnian: Timanites Genozone UD I-C. Re-gional Zone of Koenenites beckeri (17b). Conodont transitansZone; MN Zone 4.

Genus ACANTHOCLYMENIA Hyatt, 1900Type species.–Clymenia (Cyrtoclymenia) Neapolitana

Clarke, 1892, by original designation (Hyatt, 1900: 548).Diagnosis.–Acanthoclymeniids in which Manticoceras-

type suture is developed at least by 10 mm diameter. Conchsubevolute, with flattened venter and ventrolateral furrowsand ornamented in early five volutions with distinctive con-cave lateral flares on flanks. Growth lines biconvex.

Included species.–Acanthoclymenia neapolitana (Clarke,1892: 57-63, text-figs 1-12; House, 1961: pl. 75, figs 1-11,text-figs 1A-D, lectotype NYSM 3625); A. genundewa(Clarke, 1897: 53, nom. nud.; 1898, 1899a, b: pl. 8, figs 1-3); A. forcipifer (G. & F. Sandberger, 1850: 81, pl. 6, figs3a-c); A. orientale (Bogoslovsky, 1969: 89, text-fig. 19, pl. 2,figs 3-5); A. planorbe (G. & F. Sandberger, 1850: 96, pl. 9,fig. 3a; lectotype of House in House & Pedder, 1963: 517).

Discussion.–The name Acanthoclymenia is employed herefor small, evolute to serpenticonic forms with tabular ventersin which the lateral saddle is high, asymmetrical, and some-what crooked but in which the saddles and lobes lack theextreme angularity of Probeloceras exemplified by the type P.lutheri, here regarded as the primitive member of the Belo-ceratidae. Prochorites is separated from Acanthoclymenia andProbeloceras by its distinctively concave venter in the adult.

For 70 years, the type species (by monotypy) was consid-ered to be a clymeniid. Clarke (1892) originally assigned itto the genus Clymenia and later Hyatt (1900: 548) createdthe genus Acanthoclymenia with it as the only species. Schin-dewolf (1955: 422) placed the genus as the sole member ofa new family, the Acanthoclymeniidae. As a member of the

Clymeniida, the form was anomalously old, as was recog-nized in the last century. In Europe, clymeniids do not ap-pear until the Platyclymenia Stufe of the Famennian. House(1961), however, was able to demonstrate from the baritictype material that the supposed dorsal siphuncle was in factthe prolongation of a middorsal lobe and that the suture wasalso typical ofManticoceras. Acanthoclymenia and the Acan-thoclymeniidae were thus regarded as synonyms of Mantic-oceras and the Gephuroceratidae, respectively.

In resurrecting the generic name for use as a genuswithin the Gephuroceratoidea, we are following the reason-able current trend to distinguish forms previously assignedto the carpet-bag genus Manticoceras, but with distinctiveshell morphology, as distinct genera. The forms included inAcanthoclymenia here all show rather more evolute shellsthan Probeloceras, and they differ also in that they achievethe Manticoceras type suture at an early stage, whereas mostspecies of Probeloceras never show it, and those that doachieve it late. It is considered that the plexus Ponticeras/Acanthoclymenia /Probeloceras is the line of evolution intothe Beloceratidae.

The relationship of Ponticeras to the probeloceratidgroup andManticoceras has been much discussed (Glenister,1958; Clausen, 1969; Bogoslovsky, 1969; Kirchgasser,1975) but was not satisfactorily resolved until precise strati-graphical information was forthcoming from sequences inthe Canning Basin (Becker et al., 1993). Most species ofPonticeras are large in the adult compared to probeloceratids,with the problem in separation arising when comparing ju-veniles. Nevertheless we take the view that Ponticeras consti-tutes a distinct group in which the external saddle and lobesare rounded at all stages and the lateral saddle is low, sym-metrical, and broad.

Distribution.–Eastern North America, Europe, Russia,andWestern Australia. New York: Penn Yan Shale (GeneseeGroup) to Cashaqua Shale (Sonyea Group; Wyoming toSteuben counties). In New York, Acanthoclymenia enters the

section as Acanthoclymenia sp. in the Penn Yan Shale justbelow the Crosby Sandstone at Keuka Lake but it can beginslightly lower at the level of the Linden Horizon at Linden,New York. A. genundewa ranges through the GenundewaLimestone and possibly into the lower West River Shale.From higher in the West River Shale, particularly in therhythm below the Bluff Point Siltstone and in the Fossil LogHorizon near the top of the unit, pyritic inner whorls ofrather more evolute forms occur that are referred to A. aff.neapolitana. Acanthoclymenia has not been found in thelower Cashaqua Shale but reappears in the upper Cashaquaas A. neapolitana, especially in the Shurtleff Septarian Hori-zon.

Range.–Frasnian: Upper Ponticeras or NeopharcicerasGenozone UD I-A to Prochorites Genozone UD I-F. NewYork: Koenenites Genozone UD I-B, Regional Zone of K.styliophilus styliophilus (16) to ProchoritesGenozone UD I-F,Regional Zone of Prochorites alveolatus (20).

Acanthoclymenia neapolitana (Clarke, 1892)Pl. 6, Figs 1-15; Text-figs 30E-H, 32B; Table 5

Clymenia (Cyrtoclymenia) Neapolitana Clarke, 1892: 57-63, text-figs 1-12; Frech, 1913: 8.

Cyrtoclymenia Neapolitana.Clarke, 1897: 54; 1898, 1899a, b: 131,text-figs 102-106, pl. 8, figs 19-25.

Acanthoclymenia neapolitana. Hyatt, 1900: 548; Schindewolf,1923: 482; 1934: 346; Miller, 1938: 189, figs 39A-I, pl. 38,figs 17-20; Schindewolf, 1955: 421; Schindewolf in Moore,1957: L40, text-figs 39a-b; House & Kirchgasser, 1993: 276,278.

Manticoceras neapolitanum. House, 1961: 472, pl. 75, figs 1-11,text-figs 1A-D; 1962: 258; Bogoslovsky, 1969: 222; Clausen,1969: 116, 146; 1971: 182; Kirchgasser, 1975: 67.Type material.–Miller (1938: 192) mentioned 21 syn-

types (excluding one he regarded as a Manticoceras) and 29hypotypes in the NYSM. These include the lectotype(NYSM 3625) designated by House (1961: 474) and other

Table 5. Biometric data for Acanthoclymenia neapolitana (Clarke, 1892) from the Shurtleff Septarian Horizon of the upper Cashaqua Shale.Note: These delicate shells do not lend themselves to normal caliper measurments and hence the figures below are approximate. See Text-fig. 32 for graphs.


NYSM 3632 12.4 ca. 3.5 5.2 4.8 0.67 0.39NYSM 3631 8.0 ca. 2.6 2.8 3.4 0.93 0.43NYSM 12171 ca. 7.7 2.4 2.3 — 1.04 —NYSM 3629 7.5 3.0 2.3 3.8 1.30 0.51NYSM 3633 6.7 2.5 2.2 3.4 1.14 0.51NYSM 12172 6.2 ca. 2.2 1.9 — 1.16 —

4.5 1.8 1.3 — 1.38 —3.4 1.4 0.89 — 1.57 —

NYSM 3634 5.6 2.5 1.9 — 1.32 —


Text-fig. 30. Sutures and cross sections of Acanthoclymenia spp. from the Genesee and Sonyea Groups in New York State. A-C, I-L. Acan-thoclymenia genundewa (Clarke, 1898). A. NYSM 13169/WTK 3001 from a nodule bed 0.3 m (1 ft) below the top of the Genundewa Lime-stone, from the top of the main falls in Beards Creek (Loc. 18/6a), Leicester, Livingston County. B. NYSM 13167, from the same localityand horizon. C. NYSM 3645; a syntype, here designated as lectotype, figured by Clarke (1898) from the Genundewa Limestone at Genun-dewa Point (Loc. 23b), Canandaigua Lake, Yates County, X 18. I. NYSM 13162, suture at ca. 6.5 mm diameter (ca. 3.25 whorls), from theGenundewa Limestone, Beard's Creek (Loc. 18/6a) X 18. J. NYSM 13166, suture at ca. 2 whorls, based on a specimen from the samelocality and horizon, X 18. K. NYSM 13164, cross section at 7.9 mm diameter from the same locality and horizon, X 5.6. L. NYSM 13170,cross section at 8.5 mm diameter, of a specimen from the Genundewa Limestone, Seneca Point Gully (Loc. 23/1), Canandaigua Lake, On-tario County, X 5.7. D. Acanthoclymenia aff. neapolitana (Clarke, 1892), NYSM 12148, cross section at ca. 9 mm diameter of a specimenfrom the Fossil Log Horizon (Williamsburgh Bed), West River Shale, Beard's Creek (Loc. 18a/4b), Leicester, Livingston County, X 7. E-H. Acanthoclymenia neapolitana. E, G. NYSM 3625, cross section and suture based on the lectotype, thought to be from the Shurtleff Sep-tarian Horizon in the Cashaqua Shale, Shurtleff ’s Gully (Loc. 41), Livingston County; suture reversed, X 7. F, H. NYSM 11264, suture andoutline based on a specimen probably from the same locality and horizon; suture reversed, X 7. Scales = 1 mm (A); 5 mm (B).



material figured by Clarke [1899a, b: pl. 8, fig. 19 (NYSM3631), fig. 20 (NYSM 3632), fig. 21 (NYSM 3628), fig. 22(NYSM 3633), and fig. 25 (NYSM 3629). All of this mate-rial is excellently preserved as baritic shell replacements.

New material.–Nine indifferently preserved baritic re-placements (including NYSM 12171 and 12172) and twocrushed pyritic specimens, all collected by WTK.

Dimensions.–See Table 5.Description.–Detailed accounts of the type material hav-

ing been given by Clarke (1899a, b), Miller (1938), andHouse (1961), no lengthy account is required here.

Species characterized by unusual varices (festoons orflares) developed during the first five whorls on flanks andnumbering 9-11 per whorl. Varices not extending onto theventer or ventrolateral shoulder unless, as Clarke remarked,they did and were subsequently resorbed before formation ofsubsequent whorl. One specimen (Pl. 6, Fig. 1) suggestingaccommodation of a later whorl to form of the flares; mostspecimens (Pl. 6, Figs. 9, 11, 13, 15) suggesting varices re-stricted to the flanks.

Discussion.–It is the presence of lateral varices and a de-pressed whorl section in the early volution that distinguishesthis species from other widely evolute forms in the CashaquaShale. The holotype ofGephyroceras holzapfeliClarke, 1898,is a pyritic internal mold, and hence does not show the shell(the festoonlike structures seen on the figure herein (Pl. 25,Fig. 1) represent areas of crushing within camarae of thephragmocone), but it is a larger species and seems more evo-lute at comparable diameters; this species is placed here as ajunior synonym of Probeloceras lutheri. Enseites sulcatum(Matern, 1931), generically separated by Becker & House(1993: 113), differs in a grooved venter shown by outerwhorls of the internal mold and in a rather more serpenti-conic form.

Distribution.–Upper Cashaqua Shale (Sonyea Group) inWyoming (Livingston and Ontario counties). The originalsyntypic material of Clarke almost certainly came only fromthe Shurtleff Septarian Horizon in Shurtleff ’s Gully (Loc.41) or Shurger’s Glen, a tributary of Kinney Creek, Livonia(Livingston County). This level is high in the CashaquaShale (Text-fig. 10). Some of the material figured by Clarkein 1898 and 1899, however, although almost certainly fromthe same level, was recorded as from “calcareous concretionsin the vicinity of Honeoye Lake (Ontario County) and tothe westward as far as Conesus Lake” (Clarke, 1899a, b:134) and “various localities in Wyoming County” (Clarke,1899a, b: 159). Almost all of this material is in the NYSM.

New baritic material is also available from the ShurtleffSeptarian Horizon at Shurtleff’s Gully (Loc. 41) (NYSM12171 and 12172; 16580-16583) and at Cottonwood PointGully (Loc. 40/13), Conesus Lake (NYSM 16584). There

are two pyritic and crushed specimens, one from the upper,dark transition shales of the Cashaqua Shale in Beards Creek(Loc. 38/10; NYSM 16585), and the other from Buck RunCreek (Livingston County; Loc. 39/22; NYSM 16586), butthese poorly preserved specimens could be Probeloceras orProchorites.

Range.–Frasnian: ProchoritesGenozone UD I-F. RegionalZone of Prochorites alveolatus (20). Conodont Zone MN 6.

Acanthoclymenia aff. neapolitana (Clarke, 1892)Text-fig. 30D

Material.—Fourteen calcitic and pyritic replacements,mostly small juveniles, from the Fossil Log Horizon of theupper West River Shale (Williamsburgh Bed of Over et al.,1999, 2003), 3.4 m below the Middlesex Shale at BeardsCreek (Loc. 18a/4b), and three from probably the samehorizon, 3.4 m below the Middlesex Shale at Chidsey PointCreek (Loc. JJ, Py-7/2), Keuka Lake.

Discussion.–The small size and indifferent preservationmakes comparison of these specimens difficult. Howeverthey are all highly evolute to serpenticonic with a corre-spondingly low rate of whorl expansion (see cross section,Text-fig. 30D, NYSM 12148, from Beards Creek). Whereseen, the lateral saddle is rather narrow and rounded. Thesefeatures suggest closer affinity to Acanthoclymenia neapoli-tana from the Cashaqua Shale than to A. genundewa or A. cf.genundewa from the Genundewa Limestone and the Ge-nundewa Pyrite and lower West River Shale.

Acanthoclymenia sp. A of Kirchgasser (1975) (NYSM13175) from the upper West River at Whetstone Brook(Loc. 26a/1), 2 mi (3.2 km) west of Honeoye (OntarioCounty), differs from A. aff. neapolitana in its higher whorlexpansion and broader lateral saddle. The stratigraphic po-sition of A. sp. A below the Middlesex Shale is unknown butis probably lower than the Fossil Log Horizon (Williams-burgh Bed) of Beards Creek and Chidsey Point (Text-fig.9).

Distribution.–Fossil Log Horizon (Williamsburgh Bed)of upper West River Shale (Genesee Group; Livingston andSteuben counties).

Range.–Frasnian: Timanites Genozone UD I-C. Re-gional Zone of Koenenites beckeri (17b). Conodont ZoneMN 4.

Acanthoclymenia genundewa (Clarke, 1898)Pl. 24, Figs 1-4; Text-figs 30A-C, K, L; Table 6

Gephyroceras? (Probeloceras?) genundewa Clarke, 1898, 1899a, b:86, pl. 8, figs 1-3.

Manticoceras genundewa (Clarke). Miller, 1938: 88-89, pl. 20, figs1-4.

Probeloceras genundewa (Clarke). House, 1962: 256-259, text-fig.3E; Kirchgasser, 1975: 77-82, pl. 2, figs 1-6, pl. 3, figs 3-4,12-13, text-figs 7B, 8B, 9A-F, table 2.

Type material.—Clarke’s three syntypes are juveniles, thelargest (NYSM 3647) approximately 8 mm in diameter. Thelectotype, selected herein, is NYSM 3645 (Clarke, 1898: pl.8, fig. 1; Miller, 1938: pl. 20, fig. 2; refigured here, Pl. 24,Fig. 4) from the Genundewa Limestone at GenundewaPoint (Loc. 23b), Canandaigua Lake (Yates County). NYSM3646 (Pl. 24, Figs 1-2) is from the same horizon and locality,but NYSM 3647 could have come from the GenundewaLimestone at Bristol (Ontario County; Miller, 1938: 89).

Material.–New collections from the Genundewa Lime-stone are: Taunton Gully (Loc. 17/8), near Leicester (Liv-ingston County), Beards Creek, Leicester (LivingstonCounty; Loc. 18/6a), Fall Brook-Dewey Hill, Geneseo (Liv-ingston County; Loc. 19/10), Seneca Point (Loc. 23/1), andGenundewa Point (Loc. 23b/4), Canandaigua Lake (On-tario and Yates counties). Mostly calcitic replacements; somebaritic and pyritic replacements. Kirchgasser (1975) re-ported on specimens from the Genundewa Limestone atBeards Creek (Loc. 18, WTK 3001) and at Seneca Point(Loc. 23, WTK 3006a).

Description.–Little can be added to Clarke’s (1898) de-scription, summarized by Miller (1938), and to the accountgiven by Kirchgasser (1975). Conch thinly discoidal andsubevolute with whorl height appearing to increase morerapidly than in Acanthoclymenia aff. neapolitana from theupper West River Shale and A. neapolitana from the upperCashaqua Shale. Flask-shaped in cross section with roundedflanks converging to weak ventrolateral furrows and tabularventer. Growth lines with strong deep saddle on the mid-flank.

Suture of the lectotype (NYSM 3645) at 4.8 mm diam-eter, illustrated by House (1962: text-fig. 3E), with broad

asymmetrical lateral saddle and broad, rounded lateral lobe;greater asymmetry and angularity of lateral lobe is apparentat approximately 6.5 mm diameter (NYSM 13162; Kirch-gasser, 1975: text-fig. 9E).

Distribution.–Lower Genundewa Limestone (GeneseeGroup; Livingston, Ontario, and Yates counties) and prob-ably Harrell Shale in West Virginia. In the Genesee Valley(Livingston County) and near Canandaigua Lake in Ontarioand Yates counties, Acanthoclymenia genundewa appears tobe restricted to the nodular lower part of the GenundewaLimestone, below the well-bedded layers that define theupper part of the unit.

From the Harrell Shale south of Landes Post Office(Grant County), West Virginia, House (1978) assigned toProbeloceras cf. genundewa a collection of 16 specimens inwhich whorl form and cross section conformed closely withAcanthoclymenia genundewa from the Genundewa Lime-stone in New York.

The stratigraphic position of the single specimen fromthe Chut River section in Timan, Russia, described and il-lustrated by House & Becker in Becker et al. (2000) as Acan-thoclymenia aff. genundewa, is unknown.

Range.–Frasnian: Koenenites Genozone UD I-B. Re-gional Zone of K. styliophilus kilfoylei (16). Conodont ZoneMN 2, ?3.

Acanthoclymenia cf. genundewa (Clarke, 1898)

Remarks.–At the NY Rte. 20 embankment (Loc. 24) andbeneath the bridge overpass at Bethany Center (GeneseeCounty; Loc. 24/2), several pyritic replacements of juvenilesfrom the basal West River Shale have whorl forms and su-tures resembling Acanthoclymenia genundewa but better ma-terial is needed for an unqualified assignment. Similarlyproblematic are several calcitic replacements from the lower

Table 6. Biometric data for Acanthoclymenia genundewa (Clarke, 1898) from the Genundewa Limestone.


NYSM 13167 11.5 3.1 5.0 3.5 0.62 0.30NYSM 16535 ca. 9.6 — ca. 5.0 2.9 — 0.30

5.1 1.5 1.9 2.1 0.79 0.413.7 ca. 1.1 1.3 1.6 0.85 0.432.7 ca. 9.0 0.90 1.1 1.00 0.41

NYSM 13162 9.1 — 4.3 2.5 — 0.27NYSM 13170 8.5 2.3 3.7 — 0.62 —NYSM 13164 7.9 2.3 3.1 2.9 0.74 0.37

5.5 1.7 1.9 2.2 0.89 0.404.0 1.3 1.4 1.7 0.93 0.432.9 1.0 0.95 1.2 1.05 0.41

NYSM 13169 4.4 1.3 1.5 1.8 0.87 0.413.2 0.95 1.1 1.4 0.86 0.44



few meters of theWest River Shale at Beards Creek, Leicester(Livingston County; Loc. 18a/1-3).

Also included in Acanthoclymenia cf. genundewa are sev-eral pyritic replacements of juveniles from the GenundewaPyrite at Keuka Lake, including one from Cornwall Gully(Loc. DD, Py-10/1), Penn Yan (Yates County), and severalfrom Sunset Point Gully (Loc. F, Py-16/2), 9.3 km (5.8 mi)southwest of Penn Yan (Yates County).

Distribution.–Genundewa Pyrite Horizon and basal andlower West River Shale (Genesee Group) in Genesee (Liv-ingston and Yates counties).

Range.–Frasnian: Timanites Genozone UD I-C. Re-gional Zone of Manticoceras contractum (17a). ConodontZone MN 3.

Acanthoclymenia sp.

Material.–Single pyritic inner whorl collected by J. Kral-ick from a concretion in the Penn Yan Shale in the interval0.30 m (1 ft) below the base of the Crosby Sandstone atSunset Point, Keuka Lake (Yates County; Loc. F, Py-16).Small crushed molds of an evolute goniatite from the shaleon top of the Linden Horizon at Linden (Genesee County;Loc. 15/8), are indeterminate but could be Acanthoclymeniasp.

Description.–Specimen NYSM 12170 with diameter of9.0 mm. Coil moderately evolute with high whorl expan-sion. Whorl width highest near umbilical wall; flanks slop-ing gently to narrow tabular venter; distinct but shallowventrolateral furrow. Suture not seen but growth-line patternwell displayed and showing deep saddle across flank andstrong, narrow, ventrolateral salient typical of Acanthocly-menia.

Discussion.–Close affinity to Acanthoclymenia genundewafrom the Genundewa Limestone is suggested.

Distribution:–Upper Penn Yan Shale (Genesee Group)in Yates County; ?Genesee County.

Range.–Frasnian: Koenenites Genozone UD I-B. Re-gional Zone of K. styliophilus styliophilus (16). ConodontZone MN 2.

Genus CHUTOCERAS Becker & House, 2000Type species.—Chutoceras manticoides Becker & House,

2000, by original designation (Becker &House in Becker etal., 2000: 83).

Diagnosis.–Large laterally compressed acanthoclymenids,with wide umbilicus, similar to Ponticeras but not develop-ing tabular venter and with more advanced sutures compris-ing sharp lobes and rounded saddles, often rather equal anddistinguished by sharp umbilical lobe centered on or justdorsal of seam.

Included species.–The type species, Chutoceras manticoidesBecker & House (in Becker et al., 2000: 83, text-figs 8A-B,pl. 1, figs 1-2) and C. nundaium (Hall, 1874), describedherein.

Discussion.–The distinguishing feature is the ponticeratidform with a suture of the complexity of Koenenites, that ismore advanced even thanManticoceras in possessing an um-bilical lobe centered on or near the seam. The shell form in-dicates relation to Ponticeras. The type species shows atwisted first lateral lobe, a larger lateral saddle and sharp lat-eral lobe, and has the characteristic umbilical lobe probablyjust dorsad of the umbilical seam.

Distribution.–Eastern North America: Ithaca Shale andSandstone (Genesee Group) in Tompkins County; Timan,Russia.

Range.–Ponticeras or Neopharciceras Genozone UD I-A(New York): ?SandbergerocerasGenozone UD I-D, ?Probelo-ceras Genozone UD I-E (Timan, Russia).

Chutoceras nundaium (Hall, 1874)Pl. 9, Fig. 1; Text-figs 28H-K

Goniatites (Clymenia?) nundaiaHall, 1874: 3 (134-135); 1876: pl.70, figs 13, 15 (not fig. 14 = Ponticeras perlatum)

Goniatites sinuosus Hall, 1879: 460-463, pl. 70, figs 13, 15, (notfig. 14 = P. perlatum), pl. 74, fig. 11; 1888: 40, pl. 128, figs 1-2.

Ponticeras cf. regale (Holzapfel, 1899). Kirchgasser, 1985: 231-232,fig. 4b; House & Kirchgasser, 1993: 275-276.

Material.–Lectotype here designated as NYSM 3751,figured by Hall (1879: pl. 70, fig. 15, pl. 74, fig. 11). Thereare four specimens from the Ithaca Shale and Sandstonehoused in the NYSM that are assigned to this species:NYSM 3749 and 3751, originally described by Hall (1874,a valid designation) as Goniatites nundaia and later figuredand described (Hall 1876, 1879) as Goniatites sinuosus. Thelocalities given by Hall were “Lower Chemung group orIthaca Beds of the Chemung Group . . . at the inclined-planeof the railroad” at or near Ithaca (Tompkins County). Theinclined plane locality (NYSM 3749 and 3751) is at SouthHill, Ithaca (Loc. AA, Dy-10c), and the horizon is in theIthaca Shale and Sandstone at approximately the level of theUniversity Quarry at Fall Creek, Ithaca (see below). A thirdspecimen (NYSM 3750) figured by Hall (1876: pl. 70, fig.14; 1879: pl. 70, fig. 14) from “the Ithaca beds of theChemung group near Truxton, New York (CortlandCounty)“ is believed to be a Chutoceras but it is too poorlypreserved for certainty.

Two additional specimens (NYSM 3753 and 3754) weredescribed and figured by Hall (1888: 40, pl. 128, figs 1-2)as Goniatites sinuosus from the “Portage group. Ithaca,


Tompkins County.” Labels with the specimens and the mu-seum catalog (Clarke & Reudemann, 1903: 605) indicatethat they were collected by S. G.Williams from the “Portage(Ithaca beds) at the University quarry,” Ithaca, New York.There were many quarries on and in the vinicity of the Cor-nell University campus during the latter part of the nine-teenth century, but the one called the University Quarry waslocated at the edge of the Fall Creek (Loc. X, Dy-10) and theinterval of shelly limestone horizons in the Ithaca Shale andSandstone (“Firestone bed” facies of Williams, 1884: 17)that yielded the specimens is estimated to be approximately72 m (235 ft) above the base of the Renwick Shale (Kirch-gasser, 1985).

Some additional but poorly preserved specimens(crushed molds) from the Ithaca Shale and Sandstone in theIthaca area are here assigned to Chutoceras with question:CU 10928 and 10938 from approximately the level of theWilliams Brook Coquinite, approximately 53 m (175 ft)above the Renwick Shale at Williams Brook (Loc. R, I-2);CU 10943, from a quarry in the lower Ithaca, 0.5 mi (0.8km) south of Williams Brook (Loc. S, I-2a); an unlabeledCU specimen from the Ithaca in Enfield Glen (Loc. BB, I-8a); and CU 10923a from the Ithaca at Cascadilla Creek(Loc. Z, Dy-10b). A fragment with growth lines from theupper Penn Yan Shale at Mill Creek (Loc. L, Bed Ov-10/2)near Lodi (Seneca County), could represent C. nundaium.

Dimensions.–NYSM 3753, D = 110 mm, WH = 43.7mm, UW = 32.8 mm.

Diagnosis.–Large, evolute, widely umbilicate, com-pressed shells; diameter of NYSM 3753 exceeding 100 mm.Whorl-form similar to that of Ponticeras perlatum but flanksmore parallel-sided, with venter broader and flatter, andwith sharper break at ventrolateral margin. Weak ventrolat-eral furrow as in P. perlatum not seen.

Suture characterized by broad, prominent ventrolateral,lateral, and umbilical saddles that are more symmetrical andequally sized than in Ponticeras perlatum. Lateral lobes nar-rower, more angular, and pointed than in P. perlatum. Chuto-ceras manticoides differs in having umbilical lobe notcentered on seam and slightly twisted, sharp first lateral lobe.

Growth lines follow pattern seen in Ponticeras perlatumwith broad lobe centered on mid-flank leading to broad andprominent salient centered outside ventrolateral margin, andtongue-like lobe across venter.

Discussion.–When James Hall described the species Go-niatites sinuosus in 1843, only a few fragments were known,one of which is the syntype [Hall, 1843: 243, fig. 106(6)]from Cashaqua Creek (Cashaqua Shale of the Sonyea Groupat Cashaqua Creek, Sonyea, Livingston County); the spec-imen is AMNH 5887/1 (Miller, 1938: 115).

Although the name Goniatites sinuosus was widely em-

ployed for specimens subsequently discovered in what arenow the Genesee, Sonyea, and West Falls Groups (Hall,1876, 1879), Hall introduced the names Goniatites (Cly-menia?) nundaia, G. pattersoni (Hall, 1860), and G. com-planatus perlatus to differentiate some forms as speciesseparate fromG. sinuosus. Both Clarke (1898, 1899a, b) andMiller (1938: 107) concluded that G. pattersoni is a syn-onym of Manticoceras sinuosum, an opinion with which weagree. Similarly, Hall (1879) and Clarke (1898, 1899a, b)concluded that the type specimens of G. (C.?) nundaia werealso conspecific with the type of G. sinuosus and Miller(1938: 106-107) concurred. The figured types of G. (C.?)nundaia (NYSM 3749 and 3751), however, which are herereferred to Chutoceras nundaium, are clearly not M. sinuo-sum, a species characteristic of the Cashaqua Shale of theSonyea Group.

The name “nundaia“ is available for NYSM 3749 and3751. The first of these is strongly distorted and could be aPonticeras because it has a rounded lateral lobe and no lobeon or near the umbilical seam. The two specimens (NYSM3753 and 3754) assigned to Goniatites sinuosus by Hall(1888) are here assigned to Chutoceras nundaium.

Hall’s specific name, which has clear priority, might bederived from “Nundawao,” the “Great Hill people” of theIroquois whose settlement by the same name, said to be theoldest Seneca village, was located just north of Naples at thesouthern end of Canandaigua Lake (Luther, 1910). Thegroup’s first council fire took place to the north at “Genun-dewa” or “Bare Hill” on the eastern side of the lake, an areathat would link the name appropriately to rocks of theGenesee Group. However, the name also suggests the area ofthe Nunda Sandstone, a unit in the West Falls Group,named for Nunda (Livingston County), which is perhapsconfusing for specimens restricted to the stratigraphicallymuch lower Genesee Group inTompkins County, but Hall’sname has clear priority.

We have referred this group in the past to Ponticeras andto the species P. cf. regale (see Kirchgasser & House, 1981;Kirchgasser, 1985; House & Kirchgasser, 1993) to draw at-tention to their similarity to one member of a group of pon-ticeratids described by Holzapfel (1899) from the LowerDomanik Suite or Group, at localities along the Chut River,near Ukhta, in South Timan, Russia. These Timan species,redescribed by Bogoslovsky (1969), are similar to the NewYork specimens in their relatively large evolute shells andprominent growth lines with strong ventrolateral salients,but do not show the sutures with broadly rounded, symmet-rical lateral saddles, sharp lateral lobes. or umbilical lobescentred on the seams. The Lower Domanik Ponticerasspecies and Chutoceras manticoides are reported at levels withconodonts of Zone MN 5 (punctata Zone) that is much

higher than Ponticeras and Chutoceras in New York (Beckeret al., 2000).

Among the crushed molds from lower Ithaca Sandstoneand Shale that are assigned with question to Chutocerasnundaium, CU 10938 from Williams Brook and the unla-beled specimen from Enfield Glen, in particular, show clearevidence of the sharp break at the ventrolateral margin thatis characteristic of the species.

Distribution.–Ithaca Shale and Sandstone (GeneseeGroup) inTompkins County; ?upper Penn Yan Shale (Gene-see Group) in Seneca County. Chutoceras nundaium is re-stricted to the Ithaca Shale and Sandstone of the GeneseeGroup in Tompkins County, in the interval between theWilliams Brook Coquinite (and possibly lower in the Ithaca)and the Quarry Sandstone interval at Ithaca (Kirchgasser,1985). The species could be represented in the equivalentupper Penn Yan Shale in Seneca County.

Range.–Frasnian: Ponticeras orNeopharcicerasGenozoneUD I-A. Regional Zone of Chutoceras nundaium (15c).Conodont Zone MN 1 (Kirchgasser, 1994; conodonts re-covered from limestone matrix with NYSM 3753).

Genus PROCHORITES Clausen, 1969Type species.–Probeloceras alveolatum (Glenister, 1958),

by original designation (Clausen 1969: 116).Diagnosis.–Acanthoclymenids similar to Acanthoclymenia

but with grooved venter on mold and bicarinate venter onconch; suture as for family but with narrower lateral saddlewith sigmoidal ventrad face. Growth lines prorsiradiate withshallow ventral sinus and broad salient.

Distribution.–Eastern North America (Sonyea Group inNew York) and Canning Basin, Western Australia.

Range.–Frasnian: Prochorites Genozone UD I-F. Con-odont Zone MN 6, 7.

Prochorites alveolatus (Glenister, 1958)Pl. 7, Figs 4-6; Text-figs 31, 32C; Table 7

Probeloceras lutheri (Clarke). Clarke, 1898 (pars): 89-102, pl. 7,

figs 1-5 (not figs 6-10 = P. lutheri), text-figs 70-71, 73-74, 77(not text-figs 68-69, 72, 75-76 = P. lutheri).

Probeloceras lutheri (Clarke). Miller, 1938 (pars): 63-70, figs 10C-G, I (not figs 10A-B, H, J = P. lutheri), pl. 12, figs 2-3, 5-7(not figs 1, 4, 8-10 = P. lutheri).

Probeloceras alveolatum Glenister, 1958: 69-70, pl. 15, figs 2-3,text-figs 3A-B; House, 1962: pl. 45, figs 5-6, text-figs 3C-D.

Eidoprobeloceras strix Kirchgasser, 1968: 4175, nom. nud.Probeloceras strix Kirchgasser,1975: 83-85, pl. 1, figs 6-12, 15, pl.

3, figs 14-20, text-figs 6, 7B, 8B, 10, table 3.Prochorites alveolatus. House & Kirchgasser, 1993: 276.

Type specimen.–By monotypy, BMR 1727 from 187-213ft (57-65 m) above the base of the Gogo Formation, LocalityK Dup 126, approximately 4 mi southwest of Old BohemiaHomestead (126-06E;18-42S),Western Australia. Specimendescribed by Glenister (1958).

Material.–New York material was listed by Kirchgasser(1975). In addition there is a fine small specimen acces-sioned under NYSM 3760 (12306/12).

Dimensions:–See Table 7 and Text-fig. 32C for graphs.Description.–A full description of New York material

under the name Probeloceras strix was given by Kirchgasser(1975).

Remarks.–Kirchgasser (1975) removed from Probeloceraslutheri six of Clarke’s syntypes (NYSM 4061-4064, 4070,and 4071) and redescribed them with 11 additional speci-mens as P. strix. The name “strix” referred to the distinctiveconcave venter in contrast to the tabular venter in P. lutheri.The striking similarity of the New York form with the thensingle known specimen of Prochorites alveolatus from the theCanning Basin of Australia was noted (Kirchgasser, 1975:85) and subsequent discoveries of the form in Australia havedemonstrated their identity (Becker et al., 1993).

Distribution.–Upper Cashaqua Shale (Sonyea Group) inLivingston and Ontario counties, and Canning Basin,West-ern Australia.

Range.–Frasnian: ProchoritesGenozone UD I-F. Interna-tional and Regional Zone of Prochorites alveolatus (20). Con-odont Zone MN 6.

Table 7. Biometric data for Prochorites alveolatus (Glenister, 1958) from the upper Cashaqua Shale. See Text-fig. 32 for graph.


NYSM 13159 19.9 — 8.9 5.5 — 0.28NYSM 4063 17.5 4.1 8.0 4.7 0.51 0.27NYSM 13158 ca. 17.3 4.0 8.0 4.5 0.50 0.26

11.2 ca. 2.3 4.7 3.6 0.49 0.327.4 2.0 2.9 2.7 0.69 0.362.1 ca. 0.65 0.65 0.82 1.00 0.39

NYSM 13157 ca. 16.7 — ca. 7.5 ca. 4.4 — 0.26NYSM 4064 15.6 4.2 7.6 3.8 0.55 0.24NYSM 12747 4.6 — 1.8 1.8 — 0.38


Prochorites aff. alveolatus (Glenister, 1958)

Probeloceras aff. strix Kirchgasser 1975: 85, pl. 3, fig. 5.

Remarks.—A single calcitic replacement (NYSM 13176)is known from the Fossil Log Horizon, 12.8 m (42 ft) abovethe base of the Rhinestreet Shale at Buck Run Creek (Loc.65/1), Mt. Morris (Livingston County). Only the side ofthe specimen is seen; the venter and suture are notpreserved. The moderately evolute coil, thinly discoidalwhorl form with high whorl expansion and the relatively

deep umbilicus suggest affinity with Prochorites alveolatusfrom the upper Cashaqua Shale.

Distribution.–Lower Rhinestreet Shale (West FallsGroup) in Livingston County.

Range.–Frasnian: ?MesobelocerasGenozone UD I-?G. Re-gional Zone Uncertain (?21). Conodont Zone MN 7.

Family KOENENITIDAE Becker & House, 1993The family Koenenitidae comprises two stocks, one contain-ing Koenenites Wedekind, 1913, Hoeninghausia Gürich,

Text-fig. 31. Sutures and cross sections of Prochorites alveolatus (Glenister, 1958) from the Cashaqua Shale of New York State. A. NYSM4063, cross section and suture at 17.3 mm diameter based on a specimen figured by Clarke (1898: pl. 7, fig. 4) recorded as from the NaplesBeds, Honeoye Lake, and probably from the Shurtleff Septarian Horizon, Livingston County (fromHouse, 1962), X 5.2. B. NYSM 12747,suture at ca. 3.0 mm diameter of a specimen from Little Tonawanda Creek (Loc. 35/18), Linden, Genesee County, X 15. C-D. NYSM 13160,suture and cross section at ca. 5.7 mm whorl height, estimated diameter 13 mm, based on a loose specimen thought to be from the ShurtleffSeptarian Horizon,Whetstone Brook (Loc.42/WTK loc. 3004), Honeoye, Ontario County, X 5.6. E. NYSM 13159, reversed suture at 6.00mm whorl height, based on a loose specimen thought to be from the Shurtleff Septarian Horizon, Whetstone Brook (Loc.42/WTK loc.3004), Honeoye, Ontario County, X 5.7. F. Cross section after Clarke (1898: text-fig. 77) of a specimen apparently lost. Clarke did not giveeither magnification or locality; also reproduced by Miller (1938: text-fig. 10G), Miller & Furnish (1957: text-fig. 31A), and Kirchgasser(1975: text-fig. 10F). G-H. NYSM 13158, reversed suture and cross section at ca. 17.3 mm diameter based on a loose specimen thoughtto be from the Shurtleff Septarian Horizon, Whetstone Brook (Loc.42/WTK loc. 3004), Honeoye, Ontario County, X 5.6.


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Procho

ritesspp.,basedon

datagiveninTables4,5,and7.A.P

ontic

eras

perlatum

(Hall,1874)from

theLodiLimestone.B

.Aca

ntho

clym

enia

neap

olita

na(Clarke,1892)from

theShurtleffSeptarianHorizon

oftheupperCashaquaShale.C.P

roch

orite

salveolatu

s(Glenister,1958)

from

theupperCashaquaShale.


1896: 348, and Komioceras Bogoslovsky, 1958: 115, and theother stock including TimanitesMojsisovics, 1882. Only thefirst is represented in New York, but Timanites ocurs inWestern Canada (Miller, 1938).

Distribution.–North America, Europe, Russia, andWest-ern Australia.

Range.–Frasnian: Koenenites Genozone UD I-B toTimanites Genozone UD I-C, ?Beloceras Genozone I-H.

Genus KOENENITESWedekind, 1913Type species.–Goniatites lamellosus G. & F. Sandberger,

1851: 85; 1850: pl. 8, fig. 1, by subsequent designation(Wedekind, 1917: 126).

Diagnosis.–Conch subdiscoidal to stout, moderately evo-lute, with ventrolateral furrows and rounded to flattenedventer. External suture similar to that of Manticoceras butwith lateral saddle less prominent and with additional um-bilical lobe (U2). Ventrolateral lobe (E1) developing onflanks of external or ventral lobe following pattern of phar-ciceratids. Growth lines characterized by raised lirae that areprominently developed, particularly as a salient on marginsof ventrolateral furrows. Ventral wrinkle layer consisting ofdiscontinuous, wavy, closely spaced lines. Hoeninghausiasimilar but developing oxyconic venter.

Included species.–Koenenites lamellosus (G. & F. Sand-

berger, 1850), K. lamellosus kirchgasseri House, 1978, K.cooperiMiller, 1938, K.? fasciculatus (Clarke, 1898), K. hoen-inghausia (von Buch, 1832), K. styliophilus (Clarke, 1898),and K. beckeri n. sp.

Discussion.–Early representatives have broad whorl formswith flanks sloping steeply to a narrowly rounded venter,low, rounded saddles and lobes, and widely spaced growth-line lirae. By comparison, later forms have more compressed,flat-sided whorls with broadly rounded to flattened venters,sutures with high, asymmetrical lateral saddles, narrowlyrounded to angular lobes, and more closely spaced growth-line lirae.

Distribution.–North America (Squaw Bay Limestone,Michigan; Genesee Group, New York; Harrell Shale, Penn-sylvania andWest Virginia), Europe, andWestern Australia.The earliest representatives of Koenenites in New York are K.styliophilus styliophilus from the upper Penn Yan Shale and K.styliophilus kilfoylei n. ssp. from the lower Genundewa Lime-stone, both closely similar to K. cooperi from the Squaw BayLimestone of Michigan. Koenenites sp. of House (1978) andK. lamellosus kirchgasseri from the Harrell Shale ofWest Vir-ginia occur at a level equivalent to the Genundewa Lime-stone of New York. Also assigned to Koenenites, but withquestion, is K.? fasciculatus from the lower GenundewaLimestone and K.? cf. fasciculatus from the middle and upper

Table 8. Biometric data for Koenenites styliophilus styliophilus (Clarke, 1898) from the Penn Yan Shale. See Text-fig. 35 A for graph.


NYSM 3783 ca. 44 12.5 ca.18.7 10.5 0.67 0.24(lectotype) ca. 18.3 7.0 ca. 8.8 4.0 0.80 0.22

ca. 11.5 c. 4.4 ca. 5.4 ca. 2.8 0.81 0.24ca. 7.9 c. 2.9 ca. 3.4 2.0 0.85 0.25ca. 5.3 2.2 2.6 1.4 0.85 0.263.4 1.7 ca. 1.3 1.2 1.31 0.352.6 1.2 0.9 0.8 1.33 0.311.9 0.95 0.68 0.73 1.40 0.38

NYSM 12140 ca. 55 — 25.9 13.5 — 0.25NYSM 12139 ca. 38 10.2 18.1 — 0.56 —NYSM 12158 ca. 36 ca. 10.2 17.5 8.6 0.58 0.24NYSM 12159 28.5 13.3 8.5 6.9 0.64 0.24NYSM 12137 16 6.0 7.8 4.8 0.77 0.30

10.3 4.3 4.9 2.4 0.88 0.23NYSM 12138 10.1 4.6 4.6 2.6 1.00 0.26NYSM 12160 6.9 2.9 3.1 1.8 0.94 0.26

4.5 2.1 1.9 1.4 1.11 0.313.1 1.4 1.3 1.0 1.08 0.322.2 1.1 0.83 0.7 1.33 0.321.6 0.75 0.63 0.45 1.19 0.281.1 0.65 0.50 0.43 1.30 0.39

NYSM 12161 6.8 3.1 3.1 1.8 1.00 0.264.6 2.2 2.0 1.3 1.10 0.283.2 1.6 1.3 1.1 1.23 0.342.3 — 0.85 0.77 — 0.331.7 0.9 0.63 0.53 1.43 0.31



West River Shale. The last representatives of the genus inNew York are specimens assigned to K. beckeri n. sp. fromthe middle and upper West River Shale, which most closelyresemble K. lamellosus and K. hoeninghausia from Germanyand K. cf. lamellosus and K. lamellosus from the MontagneNoire, France. Species of Koenenites matching the sequencein the Penn Yan, Genundewa, and West River of New Yorkoccur with equivalent zone conodonts in the Harrell Shaleof Central Pennsylvania (Milesburg-Unionville, CenterCounty) (Kirchgasser, 1996b).

Range.–Koenenites Genozone UD I-B to TimanitesGenozone UD I-C. Conodont Zones MN 2-4.

Koenenites styliophilus styliophilus (Clarke, 1898)Pl. 8, Figs 1-4, 6, 9; Text-figs 34A-F, I-J, L, 35A; Table 8

Manticoceras pattersoni var. styliophilum Clarke, 1898, 1899a, b:47-48, 75, 82, text-fig. 55, pl. 6, fig. 30 (not text-fig. 2= K.styliophilus kilfoylei n. ssp.).

Manticoceras styliophilum Clarke. Miller, 1938: 119-121, fig. 24B(not fig. 24A = K. styliophilus kilfoylei n. ssp.), pl. 20, fig. 5.

Type specimen.–Lectotype designated herein is NYSM3783, the surviving syntype (Clarke, 1899: pl. 6, fig. 30;?text-fig. 55), from uncertain horizon and locality but prob-ably from the upper Penn Yan Shale in the region of Naples(Ontario County).

New material.–Nearly 300 specimens from 14 localitiesin the upper Penn Yan Shale and equivalents in the middleIthaca Shale and Sandstone. The majority are fragments ofmolds or calcitic replacements; some are pyritic or bariticreplacements. Included are 17 specimens in the NYSM fromthe Crosby Sandstone at Keuka Lake collected by D. Lutherin 1904 (his locality 3397). Mentioned and figured materialincludes NYSM 3783, 12134-12142, 12158-12161, and16571.

Dimensions.–See Table 8.Description.–Moderately evolute; UW/D between 0.4

and 0.3 mm to approximately 6 mm diameter, decreasing toapproximately 0.24 at higher diameters (Text-fig. 35A; Table8). Whorl form subcircular (WW/WH=1) to 7-8 mm di-ameter (3-4 volutions) becoming broadly subtriangular be-tween 10 and 20 mm diameter. Maximum whorl width(WW) at top of steep umbilical wall with flanks convergingto broadly rounded venter (Text-figs 34A-E). Beyond 20mm diameter, umbilical wall becoming less steep, flanksmore flat-sided, and venter subtabular (Text-figs 34A-B).Largest specimen observed (NYSM 12140) approximately55 mm diameter with body-whorl length of at least one-halfvolution.

A few specimens appear to be considerably more de-pressed and broader in whorl form than typical forms (Text-

fig. 34E). These somewhat more massive shells have onlybeen seen in oblique cross section and better preserved ma-terial will be needed to determine whether they are a sepa-rate morphotype or species.

Protoconch oval; diameter 0.68 mm (n = 13; range 0.58-0.80 mm); width 0.63 mm (n = 10; range 0.58-0.70 mm).Weak constriction between 0.75 and 1.0 volutions. Suturewith broad, rounded primary lobes and saddles through twovolutions. External or ventral lobe subdividing (E1-Em-E1)between 2 and 2.75 volutions (2.5-4.5 mm diameter) withEm arising in flank of ventral lobe in “pharciceratid” style(Bensaïd, 1974) rather than “manticoceratid” style, in whichE1 lobe arises from lengthening of Em saddle. Shallow inter-nal U2 lobe seen at 8 mm diameter (3.5 volutions).

Diagnostic external umbilical lobe (U2) of Koenenitesnormally seen between 15 and 20 mm diameter (fourth andfifth volutions) but delayed or incipient in some specimens(not seen at 20 mm diameter in NYSM 16571). Lateral sad-dle relatively large and asymmetrical, with steep dorsal sidebetween 20 and 30 mm diameter, but thereafter becomingmore symmetrical and distinctly flat-topped (Text-fig. 34F)as lateral lobe becomes narrowly rounded to acute.

Growth lines and ornament distinctive. Swellings ornodes developing on flanks near end of first volution, be-coming prominent in second volution, weakening throughthird and becoming reduced to narrow ribs or ridges. Diag-nostic raised lirae of the genus Koenenites arising from ridgecrests and asymmetrical with steep aboral faces. Bands offiner, normal growth lines between lirae. Pattern of raisedlirae and growth lines showing broad salient centered lowon flanks and moderately deep saddle centered dorsad ofmid-flank. Strong, narrowly curved salient of raised lirae atventrolateral margin, seen as crescent-shaped ribs in internalmolds (Pl. 8, Fig. 3). Growth lines and lirae crossing venterin high, rounded saddle.

Dorsal wrinkle layer, seen in baritized specimens, con-sisting of closely spaced, wavy lines in pattern similar to thatinManticoceras (House, 1971: pl. 1, figs 1-2). Ventral wrin-kle layer consisting of closely spaced, continuous, parallel,but moderately wavy lirae that slant gently backward acrossumbilical wall becoming rectiradiate across flanks and ven-ter.

Discussion.–Clarke regardedManticoceras styliophilum asthe “. . . characteristic variation of the specific type [Mantic-oceras pattersoni =M. sinuosum] most abundantly developedin the Styliola Limestone [Genundewa Limestone] . . . mostfrom . . . outcrops . . . on Canandaigua Lake, and a singlerepresentative . . . in the Naples fauna.” (Clarke, 1899a, b:75). Clarke only figured two specimens, an uncataloged andnow lost specimen from the Genundewa Limestone [Clarke,1899a, b: 47, text-fig. 2; refigured herein (Text-fig. 34G) as

Text-fig. 33. Sutures and cross sections of Koenenitidae, Acanthoclymeniidae and Gephuroceratidae from the middle and upper GeneseeGroup of New York. A-E. Koenenites beckeri n. sp. from the West River Shale. A-C. Specimens from a loose concretion near the Bluff PointSiltstone (Loc. XX, Nap-8/1), in the gully 0.8 mi (1.3 km) south of Middlesex, Yates County. A. NYSM 12151, cross section and suture.B. NYSM 12152, partial cross section. C. NYSM 12150, holotype, partial cross section and sutures. D. NYSM 12153, suture of a specimenfrom a concretionary level 0.67 m (2.2 ft) below the Bluff Point (Loc. 23a/3), Seneca Point Creek, Canandaigua Lake, (continued right)


Koenenites styliophilus kilfoylei n. ssp.], and the syntype fromthe Naples fauna (NYSM 3783) (Clarke, 1899a, b: pl. 6,fig. 30, text-fig. 55; refigured herein as Pl. 8, Figs 1-2) thatis the designated lectotype. The horizon and location of thespecimen is uncertain.Written in script on the original labelattached to NYSM 3783 are the words “Gon. styliophilusNaples concr.” indicating that the specimen came from aconcretion or concretion horizon but it is not clear whether“Naples” refers to the Naples fauna, a designation thatwould include horizons within the Genesee Formation, theNaples Group or Beds or Formation of older terminologythat lie above the Genesee in the Sonyea Group, or the vil-lage of Naples in Ontario County. Two labels with the spec-imen indicate the locality as Naples beds, Naples, and in theNYSM catalog the specimen is listed as from the “Portage(Naples) beds, Naples, New York.” It is important to notethat the limits of the Styliola or Genundewa Limestone werenever precisely defined by Clarke in its type area atCanandaigua Lake, and he probably included in the StyliolaLimestone the lithologically similar styliolinid limestones ofthe upper Penn Yan Shale. Furthermore Clarke’s Naplesfauna included the fauna of the Genesee Group. We believethat the surviving syntype of M. pattersoni var. styliophilum(NYSM 3783), herein determined as K. styliophilus stylio-philus, came from the Penn Yan Shale in the region ofCanandaigua Lake. Clarke’s Genundewa representatives arethe related form, K. styliophilus kilfoylei n. ssp.

The preservation of NYSM 3783, with a thick shell wallof recrystallized black calcite and raised growth-line lirae, issimilar to the specimens described herein from styliolinidbeds in the upper Penn Yan Shale, and especially from theLinden Horizon, the Crosby Sandstone, and equivalenthorizons in the Ithaca Shale and Sandstone (Kirchgasser,1985). NYSM 3783 is remarkably similar to a specimen col-lected by J. W. Wells (CU 40101, figured herein, Pl. 8, Fig.

4) and labeled as from the Genundewa Limestone atTaunton Gully in the Genesee Valley, but that almost cer-tainly came from the Linden Horizon (Loc. 17/6) or there-abouts in the upper Penn Yan Shale (J. W. Wells, pers.comm., 1974).

NYSM 3783 also closely matches specimens of compa-rable size from the Crosby Sandstone of the upper Penn YanShale in the vinicity of Keuka Lake, including a collectionreported as Manticoceras from Sartwell Ravine (Loc. C, Py-11) by D. D. Luther in 1904. Luther’s loc. 3397 at 880 ft(268 m) elevation, 1 mi south of Penn Yan, miscorrelated asCashaqua Shale, is the Crosby Sandstone (Kirchgasser,1985). Several specimens among the 17 in the NYSM col-lection have the same distinctive shell form and growth-linepattern of NYSM 3783, and one in particular of similar di-mensions (NYSM 16563, Pl. 8, Fig. 9) shows the suturewith the unmistakable U2 umbilical lobe of Koenenites.

The lost syntype figured by Clarke (1899a, b: 47, text-fig. 2; refigured herein as Text-fig. 34G) probably came fromthe Genundewa Limestone at Canandaigua Lake and is hereincluded as Koenenites styliophilus kilfoylei n. ssp.

Rare specimens with a distinctly different sutural patternfrom that described above have been noted in the CrosbySandstone and are referred to morphotype B [e.g., NYSM16572-16574 (Loc. F, Py-16) and NYSM 12141 (Text-fig.34J)]. In morphotype B specimens, the sutures are closer to-gether and in the adult the lateral saddle is asymmetrical andnot flat-topped as in the more common morphotype A. Alsothe lateral lobe of morphotype B is more angular and liesmore ventrad of the less-steep umbilical wall. The stage ofappearance of the U2 lobe in morphotype B also varies. InNYSM 16573 it is seen at approximately 27 mm diameterbut in NYSM 16574 is not seen at 35 mm diameter. Mor-photypes A and B do not appear to differ in other featuresand it is unclear whether they are sexual dimorphs or sepa-

(Text-fig. 33 continued) Ontario County. E. K. beckeri n. sp. form D, NYSM 12154, cross section of a specimen from the Beards Creek Hori-zon (Loc. 18a/5) of the West River Shale, 2.4 m (8 ft) below the base of the Middlesex Shale, Beards Creek, Leicester, Livingston County.F-G. Koenenites styliophilus kilfoylei n. ssp. from the Genundewa Limestone. F. NYSM 12145, cross section and suture based on a specimenfrom loose blocks in an embankment below NY Rte. 20 (Loc. 24/1) at Bethany Center, Genesee County. G. NYSM 12146, cross sectionat 26 mm diameter of a specimen from Linden Falls (Loc. 15a/13), Little Tonowanda Creek, Linden, Genesee County. H. Koenenites sp.,CU 40022, suture of a specimen from theWest River Shale at Hicks Point (Loc. 29a), Canandaigua Lake, Ontario County; figured byWells(1956: fig.1, pl. 82, fig. 3). I-J. K.? fasciculatus (Clarke, 1898), NYSM 12155, sutures (mature suture at ca. 51 mm diameter), from a horizonin the equivalents of the Genundewa Limestone horizon in the Ithaca Formation, 5.2-6.1 m (17-20 ft) above the base of the Crosby Sand-stone, Cornwall Gully (Loc. Py-10/1), Penn Yan, Yates County, collected by Gordon Baird. K-L.Manticoceras lamed aff. cordatum (G. & F.Sandberger, 1851), NYSM 16568, sutures and cross section at ca. 9.3 mm diameter of a specimen from the Fossil Log Horizon (Loc.18a/4b) in the West River Shale, 3.4 m below the base of the Middlesex Shale, Beards Creek, Leicester, Livingston County. M, Ponticeras?sp., NYSM 12169, cross section at ca. 5.6 mm diameter, of a specimen from the Beards Creek Horizon (Loc. 18a/5), West River Shale, 2.4m (8 ft) below the base of the Middlesex Shale, in Beards Creek, Leicester, Livingston County, X 9.6. N. K. styliophilus kilfoylei n. ssp. fromthe Genundewa Limestone, NYSM 12147, whorl form and sutures (reversed) of a specimen from Loc.24/1, X 2.2. O. K. styliophilus kilfoylein. ssp. form A, NYSM 12144, suture at ca. 27 mm diameter of a specimen from the Genundewa Limestone at Bethany Center (Loc. 24/1),Genesee County, X 2.9. Scales = 5 mm.


Text-fig. 34. Sutures and cross sections of Koenenites spp. from the Genesee Group in New York. A-D. Koenenites styliophilus styliophilus(Clarke, 1898). A. NYSM 3783, cross section based on the lectotype at 44 mm diameter, figured by Clarke (1898: text-fig. 55), recordedas from the Naples Beds at Naples, Ontario County; thought to be from the Linden Horizon, Penn Yan Shale in the Naples region. B.NYSM 12136, cross section of a specimen at 28.5 mm diameter, from the Penn Yan Shale, the concretionary level (Loc. 17/5) immediatelybelow the Linden Horizon in Taunton Gully, Livingston County. C. NYSM 12137, cross section at 16 mm diameter of a specimen fromthe Linden Horizon (Loc. 15/8), Penn Yan Shale, Linden, Genesee County. D. NYSM 12138, cross section at 10.1 mm of a specimen fromthe Linden Horizon (Loc. 15/8), at the same locality. E. K. styliophylus styliophilus?, NYSM 12142, cross section at ca. 33 mm diameter ofa specimen from the Penn Yan Shale (Loc. 17a/3), Spezzano Gully, Livingston County. F. K. styliophilus styliophilus, NYSM 12139, sutureat 38 mm diameter and whorl form at 18.1 mm whorl height of a specimen from the Crosby Sandstone (Loc. C. Py-11), Penn Yan Shale,at Sartwell Ravine, Keuka Lake, Yates County, collected by D. D. Luther (NYSM collection from his locality 3397). (continued right)


rate species. In suture pattern, morphotype B resembles Koe-nenites styliophilus kilfoylei n. ssp. from the lower GenundewaLimestone to which it might be allied.

Koenenites styliophilus styliophilus is similar to K. cooperifrom the Squaw Bay Limestone of Michigan. In K. cooperi(see biometric data inTable 9), the adult whorl form is widerand more broadly rounded across the venter. The suturalontogenies are closely comparable but in the adult suture ofK. cooperi, the lateral saddle becomes high and narrow andstrongly asymmetrical (Miller, 1938: fig. 26B); at compara-ble stages in K. styliophilus styliophilus, the lateral saddle isbroader, more symmetrical, and distinctly flat-topped. Thesuture of K. cooperi is somewhat closer in form to morpho-type B and its suture and whorl form most closely resembleK. styliophilus kilfoylei n. ssp. from the Genundewa Lime-stone.

Compared to Koenenites styliophilus styliophilus the typesof K. lamellosus from Oberscheld, Germany, have a ratherbroader whorl form across the flanks and venter and a suturewith a higher, narrower, and U-shaped lateral saddle flankedby deeper E1 and U2 lobes with the latter being distinctly

angular. In these features, K. lamellosus more closely resem-bles K. styliophilus kilfoylei n. ssp. from the GenundewaLimestone and especially K. beckeri n. sp. from the WestRiver Shale.

Koenenites lamellosus kirchgasseri from the Harrell Shalenear Landes (Grant County), West Virginia, has a muchmore lingulate lateral saddle and U2 lobe than K. styliophilusstyliophilus and in K. sp. of House (1978), also from the Har-rell Shales near Landes. The whorl form is broader and theexternal sutural elements are more narrowly rounded than inK. styliophilus styliophilus. These West Virginia forms pre-sumably represent an earlier stage in the evolution of Koe-nenites than the Penn Yan and Genundewa representativesfrom New York.

Distribution.–Upper Penn Yan Shale and equivalents inIthaca Shale and Sandstone (Genesee Group; Wyoming toTompkins counties).

Koenenites styliophilus styliophilus is restricted to theupper Penn Yan Shale and is best represented in the LindenHorizon and surrounding styliolinid beds, the equivalent orslightly younger Crosby Sandstone, and the eastern repre-

(Text-fig. 34 continued) G. K. styliophilus kilfoylei n. ssp., tracing of whorl form by Clarke (1899: text-fig. 2) as Manticoceras pattersoni var.styliophylum and also figured by Miller (1938: text-fig. 24a). The specimen, which Clarke & Ruedemann (1903) did not list in their catalog,and which Miller did not report seeing, is apparently lost. It probably came from the Genundewa Limestone in the vicinity of CanandaiguaLake. Enlarged but scale unknown. H. K. beckeri n. sp., NYSM 12156, partial whorl form of a specimen from a loose concretion near theBluff Point Siltstone (Loc. XX, Nap-8/1) of theWest River Shale, in a gully 0.8 mi (1.3 mm) south of Middlesex, Yates County. I-J. K. stylio-philus styliophilus. I. NYSM 12140, suture at 16 mm whorl height of a specimen from Loc. C, Py-11. J. NYSM 12141, suture at 29.5 mmdiameter based on a specimen of morphotype B from the Crosby Sandstone (Loc. F, Py-16/1), Penn Yan Shale, from Sunset Point Gully,Keuka Lake (East Branch), Yates County. K. K. beckeri n. sp. form D, NYSM 12157, partial whorl form of a specimen from the BeardsCreek Horizon (Loc. 18a/5) of theWest River Shale, 2.4 m (8 ft) below the base of the Middlesex Shale, Beards Creek, Leicester, LivingstonCounty. L. Same as F. Scales = 5 mm.

Table 9. Biometric data for Koenenites cooperi Miller, 1938, from the Squaw Bay Limestone near Alpena, Michigan.


MMP 13846 ca. 72.0 ca. 17.0 31.8 ca. 18.0 0.53 0.25ca. 58.0 14.5 — 13.5 — 0.2341.1 — 21.5 — — —38.0 — 17.3 8.0 — 0.2135.4 ca. 10.8 17.8 — 0.61 —32.1 — 16.1 7.5 — 0.2331.5 — 14.3 7.8 — 0.25ca. 30.0 9.6 13.9 — 0.69 —ca. 27.5 — 12.7 6.5 — 0.24

USNM 96456a 66.2 16.0 28.7 16.0 0.56 0.24(lectotype)

USNM 96546b 31.4 9.6 14.7 7.5 0.65 0.2424.0 7.7 8.0 5.4 0.96 0.2319.4 6.4 9.1 4.2 0.70 0.2214.6 5.2 6.6 3.6 0.79 0.25

USNM 96546c 38.4 11.5 19.1 7.4 0.60 0.19USNM 96546d ca. 68.0 ca. 18.0 29.0 17.0 0.62 0.25USNM 96546f 16.0 6.0 6.8 4.6 0.88 0.29


Text-fig.35.Graphsshowingtheontogeny

ofKoene

nites,M

antic

oceras,and

Prob

elocerasspp.A.K

oene

nitessty

lioph

ilusstylio

philu

s(Clarke,1898),basedon

specimensfromthePenn

YanShale;basedon

datainTable8.B.M

antic

oceras

cont

ractum

(Clarke,1898),basedon

acotype

(NYSM

5125)fromtheGenundewaLimestone;M

.nod

ifer(Clarke,1885),based

ontype

material,NYSM

14994a,b,fromtheGenundewaLimestone;and

M.sin

uosum

appr

imatum

Clarke,1898,based

ontype

material(NYSM

3732)fromtheGenundewaLime-

stone.C.P

robe

loceraslut

heri(Clarke,1885),basedon

datafrom

materialfromtheCashaquaShaleinTable22.


sentatives or equivalents of these horizons in the Ithaca For-mation at Seneca and Cayuga lakes (Seneca and Tompkinscounties; Kirchgasser, 1985).

Range.–Frasnian: Koenenites Genozone UD I-B. Re-gional Zone of K. styliophilus styliophilus (16a). ConodontZone MN 2.

Koenenites styliophilus kilfoylei n. ssp.Pl. 8, Figs 5, 7-8, 12-13; Text-figs 33F, G, N-O, 34G; Table 10

Manticoceras pattersoni var. styliophilum Clarke, 1899: 47, text-fig.2.

Koenenites aff. styliophilus House & Kirchgasser, 1993: 276-277.

Etymology.–Named in honor of Clinton Kilfoyle, formerCurator of Paleontology at the New York State Museum.

Type specimen.–NYSM 12143 (Pl. 8, Figs 7-8), from thelower 15 cm of the Genundewa Limestone, from looseblocks at NY Rte. 20 embankment at Bethany Center(Genesee County; Loc. 24/1). The lost syntype and uncat-aloged specimen of Manticoceras pattersoni var. styliophilumfrom the Genundewa Limestone figured by Clarke (1899a,b: text-fig. 2; refigured herein, Text-fig. 34G) is included inKoenenites styliophilus kilfoylei n. ssp.

Material.–Over 100 specimens from the lower Genun-dewa Limestone: 74 from loose blocks of the lower 15 cmof the unit at the NY Rte. 20 embankment, Bethany Center(Genesee County; Loc. 24/1); 6 from Murder Creek, Gris-wold (Genesee County; Loc. 14a/3); 23 from Linden Falls,Linden (Genesee County; Loc. 15a/12); 4 from BeardsCreek, Leicester (Livingston County; Loc. 18a/6a); severalfrom the pyritic nodular horizon equivalent to the Genun-dewa near Keuka Lake: Cornwall Gully, Penn Yan (YatesCounty; Loc. DD, Py-10); Willow Grove (Yates County;

Loc. EE, Py-14); Sunset Point (Loc. FF, Py-16), creek aboveSunset Point (Yates County). Mentioned or figured materialincludes NYSM 12143-12147, and 12162-12165.

Dimensions.–See Table 10.Diagnosis.–Closely similar to nominate form but slightly

more evolute, broader in whorl form in early stage, moreflat-sided on flanks, with more broadly rounded venter andsuture with asymmetrical lateral saddle that is not flat-topped.

Description.–Shell form and suture similar to nominateforms from Penn Yan Shale but slightly more evolute andbroader in cross section in early stages (WW/WH higher)and becoming more flat-sided on flanks and more broadlyrounded across venter (Text-figs 33F-G, N, 34G). Suturesimilar to nominate form in having broadly rounded lateralsaddle. Lateral saddle compares most closely with morpho-type B of Koenenites styliophilus styliophilus in having lateralsaddle more asymmetrical and not as flat-topped as mor-photype A of K. styliophilus styliophilus (Text-figs.33N, 34J).Included here is single morphotype (form A, NYSM 12144)with unusual suture of closely spaced, tongue-shaped lateralsaddles with rounded crests (Pl. 8, Figs 12-13); lateral saddlenarrower and more symmetrical than typical K. styliophiluskilfoylei n. ssp. in lower Genundewa and not as flat-toppedas typical form of K. styliophilus from upper Penn Yan Shale.

As in Koenenites styliophilus styliophilus, stage of appear-ance of U2 lobe varies. In NYSM 12147, seen at approxi-mately 18 mm diameter (Text-fig. 33N) and in NYSM12144, at approximately 27 mm (Text-fig. 33O), but notin NYSM 12165 at approximately 26 mm diameter.Growth lines and wrinkle layer similar to K. styliophilusstyliophilus but raised growth-line lirae not as strongly de-veloped.

Table 10. Biometric data for Koenenites styliophilus kilfoylei n. ssp. from the Genundewa Limestone.


NYSM 12162 34.5 ca. 10 15 8.5 0.67 0.2523.3 ca. 8 10.5 6 0.76 0.26

NYSM 12144 ca. 27 — 12.5 ca. 8.5 — 0.31NYSM 12145 26.7 9 12 6.8 0.75 0.25

18 ca. 6.5 8.3 4.2 ca. 0.78 0.2311.6 ca. 4.8 5.6 2.7 ca. 0.86 0.23

NYSM 12146 26.0 ca. 10 11.3 7 0.88 0.2717.2 ca. 7.3 7.4 4.5 0.99 0.2611.6 5.4 5.1 3.3 1.06 0.287.8 3.4 3.3 2.5 1.03 0.325.2 2.5 2.0 2.1 1.25 0.403.8 1.6 1.3 1.7 1.23 0.45

NYSM 12163 26 9.2 11.8 6.6 0.77 0.2517 6.4 7.6 5.3 0.84 0.31

NYSM 12164 19 ca. 6.5 8.4 5.3 0.77 0.27ca. 13.3 — 5.7 4.3 — 0.32


Discussion.–Koenenites styliophilus kilfoylei n. ssp. com-pares closely with morphotype B of K. styliophilus styliophilusof the upper Penn Yan Shale and is a likely descendant ofthat taxon. Close similarity with K. cooperi of the Squaw BayLimestone, Michigan, is seen in whorl form and suture butin K. cooperi the asymmetry of the lateral saddle is strongerand the external U2 lobe more fully developed. Similarly thesuture of K. lamellosus is more advanced in having higherlateral saddles and lobes and more strongly developed U2lobe.

Distribution.–Lower Genundewa Limestone (GeneseeGroup) in Genesee and Livingston counties and equivalentpyritic horizon near Keuka Lake in Yates County and inequivalents of the Genundewa Limestone in the HarrellShale at Milesburg and Unionville (Center County), Penn-sylvania.

Range.–Frasnian: KoenenitesGenozone UD I-B RegionalZone of K. styliophilus kilfoylei (16b). Conodont Zone MN2.

Koenenites? fasciculatus (Clarke, 1898)Pl. 18, Figs 1-2, 5; Text-figs 33I-J

Manticoceras fasciculatum Clarke, 1898, 1899a, b: 71-74, 81-82,pl. 6, figs 13-22, text-figs 45-53, 63-64.

Manticoceras fasciculatum Clarke. Miller, 1938: 85-87, text-fig. 15,pl. 15, figs 1-10.

Types.–Nine syntypes of this species are housed in theNew York State Museum as NYSM 3736-3744. All are smalljuveniles of less than 5 mm diameter. According to Clarke(1898), all are from the Genundewa Limestone atCanandaigua Lake and Middlesex (Yates County). A lecto-type is here designated as NYSM 3738 (Pl. 18, Figs 1-2),from the Genundewa at Canandaigua Lake.

New material.–Eleven specimens from the GenundewaLimestone and pyritic nodular equivalent of the Genundewa(Genundewa Pyrite) near Keuka Lake (Yates County); onecalcitic replacement (NYSM 16575) of the inner whorlsfrom the Genundewa Limestone at Genundewa Point (YatesCounty; Loc. 23b/4), 0.71 m above the base of the unit;two partly pyritic crushed molds of adult specimens fromthe Genundewa Pyrite at Cornwall Gully, Penn Yan (YatesCounty; Loc. DD, Py-10); seven pyritic and one baritic re-placements of inner whorls from the Genundewa Pyrite atSunset Point Gully, Keuka Lake (Yates County; Loc. FF, Py-16).

Description.–Like syntypes, most new specimens aresmall inner whorls and consequently little can be added toClarke’s (1898) descriptions and illustrations summarizedby Miller (1938). Largest types [Clarke, 1899a, b: pl. 6, fig.19 (lost syntype) and 21 (NYSM 3738)] at four volutions

show moderately evolute coil with deep umbilicus withinwhich distinctive, paired, fasciculate growth lines are dis-played. These features clearly seen in NYSM 16575 fromGenundewa Limestone at Genundewa Point (Loc. 23b/4)and in baritic specimen (NYSM 16590) from GenundewaPyrite at Sunset Point (Loc. FF, Py-16/2).

Innermost whorls, preserved as pyritic replacements intypes and new material from Genundewa Pyrite, havestrongly depressed whorl form with broadly rounded venterand distinctive, broad, rib-like swellings on flanks (Miller,1938: fig. 15). Swellings appear to mark sites of raised, fas-ciculate growth lines seen in calcitic and baritic replace-ments.

Discussion.–The immature suture is distinguished by alarge, prominent ventral lobe from the sides of which theventrolateral lobes develop in the fourth whorl. This patternof sutural development on the venter, seen in the syntypesand new material, was recognized as primitive by Clarke(1898) and unlike the pattern seen inManticoceras in whichthe ventrolateral lobes arise from the enlargement of the E1saddles. This pharciceratid-type of evolving external lobe isalso seen in Pseudoprobeloceras, Acanthoclymenia, otherspecies of Koenenites, and probably alsoHoeninghausia (Ben-saïd, 1974). Because of their small size, the syntypes of K.?fasciculatus provide no information about the mature suture.One of the specimens (NYSM 12155), however, from theGenundewa Pyrite at Cornwall Gully (Loc. Py-10/1) showsthe suture of a Koenenites at approximately 51 mm diameterwith a deep lateral lobe, high saddle across the umbilicalwall, and a clearly defined U2 lobe outside the seam (Text-fig. 33J). The pyritic inner whorls of the specimen are poorlypreserved and although the irregular wrinkle-layer patternof Koenenites is seen, the typical rib-like swellings of Clarke’sspecies Manticoceras fasciculatum are not seen. Thus thegeneric assignment of K.? fasciculatus and the assignment ofNYSM 12155 to the species are problematic.

Distribution.–Lower Genundewa Limestone (GeneseeGroup), Canandaigua Lake (Yates County). Also in Genun-dewa Pyrite equivalent of the Genundewa near Keuka Lake(Yates County).

Range.–Frasnian: KoenenitesGenozone UD I-B; ?Timan-itesGenozone UD I-C. Regional Zone of K. styliophilus kil-foylei (16b), ?Manticoceras contractum (17a). ConodontZone MN 2-?3.

Koenenites? cf. fasciculatus (Clarke, 1898)

Material.–Thirty-three pyritic inner whorls recoveredfrom a loose concretion, probably from the first rhythm inthe West River Shale beneath the Bluff Point Siltstone in agully 0.8 mi (1.3 km) south of Middlesex (Yates County;


Loc. XX, Nap-8/1). One pyritic replacement of an innerwhorl from the Fossil-Log concretion horizon in the upperWest River Shale (Williamsburg Bed of Over et al., 1999,2003), 3.4 m below the Middlesex Shale in Beards Creek,Leicester (Livingston County; Loc. 18a/4b).

Discussion.–The quality of preservation varies widely butall of the specimens conform closely in whorl form and su-ture to Clarke’s types of Koenenites? fasciculatus and the newspecimens assigned to that species from the lower Genun-dewa Limestone and equivalent Genundewa Pyrite in thevinicity of Keuka Lake (Yates County). It is uncertainwhether these inner whorls from the middle and upperWestRiver Shale are the same as K.? fasciculatus from the Genun-dewa Limestone or whether they are early stages of the largeform from the same horizons referred here to K. beckeri n.sp.

Distribution.–Middle and upper West River Shale(Genesee Group) in Livingston and Yates counties.


Koenenites beckeri n. sp.Pl. 8, Figs 10-11, 14; Text-figs 33A-E, 34H, K

Type material.–Holotype here designated, NYSM 12150(Pl. 8, Fig. 14; Text-fig. 33C) from the rhythm below theBluff Point Siltstone near Middlesex (Loc. XX, Nap-8 of deWitt & Colton, 1978).

Etymology.–Named in honor of our colleague, R. T.Becker, University of Münster, Germany.

Previously referred to Koenenites sp. C (Kirchgasser &House, 1981: 42, text-fig. 2) and K. aff. lamellosus (seeHouse & Kirchgasser, 1993: 277, fig. 7).

Additional material.–Twelve specimens from the rhythmbelow the Bluff Point Siltstone of the middle West RiverShale and the correlative horizon in the Ithaca Shale andSandstone and two specimens from the upper West RiverShale; three calcitic replacements/molds (including NYSM12153) from the gray shale in the first rhythm beneath theBluff Point Siltstone (0.76 m below the horizon) at SenecaPoint Creek, Canandaigua Lake (Ontario County; Loc.23a/3); six calcitic replacements (including the holotypeNYSM 12150, and 12149, 12151-12153, 12156) from aloose concretion probably from the same first rhythm be-neath the Bluff Point Siltstone in a gully 0.8 mi (1.3 km)south of Middlesex (Yates County; Loc. XX, Nap-8/1); twomolds from a dark gray shale beneath the horizon of theBluff Point Siltstone at 920 ft (280 m) elevation and 24 mbelow the Middlesex Shale in Mill Creek, near Lodi (SenecaCounty; Loc. LL, OV-10/16); one mold from dark gray

shales beneath a black shale near the level of the Bluff PointSiltstone at 1,030 ft (314 m) elevation and 30 m below theMiddlesex in Sheldrake Creek, Cayuga Lake (SenecaCounty; Loc. MM, Gen-1a); one mold in a silty shale in theIthaca Shale and Sandstone near the level of the Bluff PointSiltstone at approximately 980 ft (299 m) elevation and 47m below the Middlesex in Lick Brook, Ithaca (TompkinsCounty; Loc. OO, I-8); two molds from a gray shale 4.9 mbelow the Middlesex Shale in Beards Creek, Leicester (Liv-ingston County, Loc. 18a/4a); one calcite and barite replace-ment from a concretion horizon 5 m below the Middlesexin Snyder’s Gully, Woodville, Canandaigua Lake (OntarioCounty; Loc. 46/c1).

Diagnosis.–Closely similar to Koenenites lamellosus butwith more flat-sided to nearly parallel-sided flanks andbroader to nearly flat venter. Later saddle of K. beckeri n. sp.is slightly more angular than in K. lamellosus.

Description.–Shell form of large specimens moderatelyevolute with flattened and nearly parallel flanks that slopegently to broadly rounded to flattened venter. Somewhatrectangular cross section of flanks and venter in largest spec-imens is distinctive.

Adult suture with high, strongly asymmetrical lateralsaddle, narrowly rounded to angular lateral lobe, relativelyhigh saddle on umbilical wall, and shallow U2 lobe on orjust outside umbilical seam. Growth lines with raised liraetypical of Koenenites that, where seen on inner whorls, aresomewhat fasciculate. On flanks of largest specimens,growth lines show sinus on flank and strong tongue-like ven-trolateral salient and broadly rounded sinus across venter.

Discussion.–The whorl form and suture indicate an affin-ity with Koenenites lamellosus. In the types of K. lamellosusfrom Oberscheld, Germany, however, the flanks are not asflat-sided and the lateral saddle is more rounded than in theNew York specimens at comparable stages.

The New York specimens are also similar to the speci-mens referred to Koenenites cf. lamellosus and K. lamellosus byHouse et al. (1985: figs 8A-C) from Bed 58 in the CPS-Esection in the Montagne Noire, France, but again they differin having flatter flanks at comparable stages. The specimenreferred to Hoeninghausia cf. archiaci from the slightlyyounger Bed 24 in Trench A’ at La Serre, in the MontagneNoire, has a similar suture (House et al., 1985: pl. 3, figs10-12, text-figs 8D-E) but at a stage twice the size of theNew York specimens the flanks slope much more steeply toa more narrowly rounded venter. As noted earlier, the genusHoeninghausia (typeH. archiaci) is restricted to undoubtedlyoxyconic forms. The specimen from Bed 24 in Trench A’ atLa Serre is a transitional form best referred to K. cf. lamel-losus.

The type specimen of Ammonites hoeninghausia from


Bensberg, Germany, which was reillustrated and assigned toHoeninghausia (Koenenites) hoeninghausia by Bensaïd (1974:107, pl. 3, fig. 7, text-fig. 14), shows a remarkable similarityto the New York specimens of comparable size in its flat-tened flanks and broadly flattened venter and acute ventrallobe, but its U2 lobe is acute as well. The horizon of vonBuch’s specimen is unknown but its suture suggests a moreadvanced member of the K. lamellosus lineage than the typeof the species and the specimens described herein as K. beck-eri n. sp. from the middle and upper West River Shale.

Distribution.–Middle and upper West River Shale(Genesee Group) in Livingston, Ontario, Yates, Seneca, andTompkins counties.


Koenenites beckeri n. sp., form DText-figs 33E, 34K

Discussion.–We previously referred to Koenenites sp. D(Kirchgasser & House, 1981: 43, text-fig. 2) and K. aff.lamellosus sp. D (House & Kirchgasser, 1993: 277, text-fig.7), two specimens (NYSM 12154 and 12157) from theBeards Creek Horizon (Bed 18a/5) in upper West RiverShale, 2.4 m below the Middlesex Shale, at Beards Creek,Leicester (Livingston County; Loc. 18a). NYSM 12157(Text-fig. 34K), which we regard as the type, is a fragmentof the outer whorl of a large individual. The raised growth-line lirae seen on the outer flank is of the Koenenites type. Incross section, the flanks slope to a broadly rounded ventergiving a distinctly different profile than the parallel-sidedprofile with broadly flattened venter seen in the specimenslower down in theWest River Shale referred to K. beckeri n.sp. Although the specimen is crushed, the cross section ofNYSM 12154 also shows flanks that are not subparallel butclearly slope toward the venter.

Assigned to the same taxon is a third specimen (NYSM16576) that shows a similar profile but a somewhat morebroadly flattened venter. Specimen NYSM 16576 is fromthe fossil log concretion horizon in the upper West RiverShale, 3.4 m below the Middlesex Shale in Chidsey PointCreek, Keuka Lake (Steuben County; Loc. KK, Py-7/2;Williamsburgh Bed of Over et al., 1999, 2003).

Distribution.–Upper West River Shale (Genesee Group)in Livingston and Steuben counties.


Family GEPHUROCERATIDAE Frech, 1897The gephuroceratids comprise the family named byWedekind (1913) as Manticoceratidae after its best knowngenus, Manticoceras, but the earlier name of Frech has pri-ority and widest use. The group is thought to have arisenfrom late Givetian serpenticonic acanthoclymeniids such asPseudoprobeloceras by a tendency toward a more involuteshell with very varied morphology, an emphasis of the mid-dorsal saddle, and the addition of umbilical lobes. Growthlines are biconvex but in some later members (Crickites) be-come convex. The family characterizes the Frasnian world-wide but became extinct at its close with the UpperKellwasser Event (Becker & House, 1993, 1994a).

The family has three named subfamilies, Gephurocerati-nae, Virginoceratinae, and Crickitinae, but the characters ofthe first group are so varied as to indicate the division israther an arbitrary one.

Included genera in the Gephuroceratinae are Mantico-ceras Hyatt, 1884, Clauseniceras Becker & House, 1993,Costamanticoceras Becker & House, 1993, DelphicerasBecker & House, 2000 (replacement for Delphinites Becker& House, 1993, non Sayn, 1901), Maternoceras Clausen,1971, Playfordites Becker et al., 1993, TrimanticocerasHousein House & Ziegler, 1977, Mixomanticoceras Becker et al.,1993, Sphaeromanticoceras Clausen, 1971, and TimanocerasBogoslovsky, 1957. The four known in New York areMan-ticoceras, Delphiceras, Playfordites, and Sphaeromanticoceras.

Included in the subfamily Virginoceratinae Yatskov,1990, are the generaVirginoceras Ruzhencev, 1960, Carino-ceras Lyashenko, 1957, and Neomanticoceras Schindewolf,1936. Of these only Carinoceras is recognized in New York.

The subfamily Crickitinae Wedekind, 1913, is charac-terized by convex growth lines from an early stage. Crickitesoccurs in New York and Canada.

In the text below, genera are treated in their approximateorder of appearance in New York.

Distribution.–Worldwide.Range.–Frasnian: Manticoceras Stufe: ?Koenenites Geno-

zone UD I-B to Crickites Genozone UD I-L. ConodontZones MN 3-13.

Subfamily GEPHUROCERATINAE Frech, 1897GenusMANTICOCERAS Hyatt, 1884

Type species.–Goniatites simulator Hall (1874: 2), bymonotypy.

Diagnosis.–Conch usually large, evolute in inner whorlsthen subevolute to subinvolute. Shell without ribs or nodes;growth lines biconvex. Suture with three pointed or roundedlobes between small ventral lobe (Em), and umbilical seam:saddle between E1 and L (Text-fig. 25) always large andhigh.


Included species.–Even after the recent separation of sev-eral genera, there are still over 40 specific names that couldbe used in this genus. Only those relevant to the descriptionof New York taxa will be referred to below. A full review ofthe genus is needed.

Discussion.–Aspects of the taxonomy of the genus werereviewed byMiller (1938: 72) and the evolutionary relationsof the group have been discussed by Becker & House(1993).We followMiller in consideringGephurocerasHyatt,1884, and Gephyroceras as junior synonyms ofManticoceras.

Distribution.–Worldwide except for polar regions andSouth America. New York: Upper Genundewa Limestone(Genesee Group) to Angola Shale (West Falls Group).Man-ticoceras first appears in New York in the Upper GenundewaLimestone with the entry ofM. contractum,M. nodifer, andM. sinuosum apprimatum, the latter two also presumablyfrom the Upper Genundewa.

Range.–TimanitesGenozone UD I-C to CrickitesGeno-zone UD I-L. Conodont Zones MN 3-13. New York:TimanitesGenozone UD I-C toNeomanticocerasGenozoneUD I-J. Conodont Zones MN 3-?11.

Manticoceras contractum Clarke, 1898Pl. 19, Figs 10-16; Text-fig. 35B

Manticoceras contractum Clarke, 1898, 1899a, b: 69-70, 81, text-figs 44, 63-64, pl. 6, figs 1-2.

Manticoceras contractum. Miller, 1938: 81-82, pl. 13, figs 12-14.

Type material.–Three cotypes of Clarke (NYSM 3734,3735, and 5125) from the Genundewa Limestone nearMiddlesex (Yates County) and Canandaigua Lake (YatesCounty). NYSM 3735 (Clarke, 1898: pl. 6, fig. 2; andherein Pl. 19, Figs 12-15), Genundewa Limestone,Canandaigua Lake, is here designated as lectotype.

New material.–Several specimens (some baritic) fromloose blocks of the Genundewa at the base of the high fallsin Fall Brook (Loc. 19a), near Geneseo (Livingston County),are assigned to this species (WK 3011, 3872); the blocks arebelieved to have come from the top of the GenundewaLimestone. Three fragments (WK 3946/1/2/7) from Bed19/10, at the top of the nodular Lower Genundewa at FallBrook-Dewey Hill (Loc. 19) are assigned with question tothis species.

Diagnosis.–Summarized fromMiller (1938) after Clarke(1898, 1899a, b): Small, subglobular conch with narrowumbilicus, rounded umbilical shoulders and ventrolateralmargin, and broadly rounded venter. Ornamentation pecu-liar with growth lines weakly biconvex and grouped into fas-cicles or bundles at maturity (Pl. 19, Fig. 14). Sutureunknown.

Remarks.–Additional material will be required to better

characterize this species. Specimen 3872/9 from the Genun-dewa at Fall Brook (Loc. 19a) has a growth-line pattern andbroadly rounded venter that matches the features of the des-ignated lectotype (NYSM 3735). NYSM 5125 at 22.3 mmhas a rather narrow whorl-form (Miller 1938: pl.13, fig. 14)and it could be a Koenenites from the upper Penn Yan Shale.

Distribution.–Upper Genundewa Limestone (GeneseeGroup) in Livingston and Yates counties.

Range.–Frasnian: Timanites Genozone UD-I-C. Re-gional Zone of Manticoceras contractum (17a). ConodontZone MN 3.

Manticoceras sinuosum apprimatum Clarke, 1898Pl. 17, Figs 5-10; Text-figs 35B, 42F

Manticoceras apprimatumClarke, 1898, 1899a, b: 62-63, 80, text-figs 63-64, pl. 6, figs 27-29; Miller, 1938: 76, pl. 13, figs 9-11;Becker & House, 1993: 119.

Type material.–The three syntypes of Clarke (NYSM3731-3733; formerly 12301/1-3) are refigured here andNYSM 3732 is here designated as lectotype.

Discussion.–Clarke’s types are poorly preserved; thus thecharacteristics of M. apprimatum will always be poorly un-derstood. The narrow umbilicus, broadly rounded whorl-form, and gently sinuous biconvex growth lines seen inNYSM 3732 suggest assignment toManticoceras but unfor-tunately the suture is not seen in any of the types. It couldwell be the earliest representative of the M. sinuosum groupand tentatively we refer to it as a subspecies of that form. Asnoted byMiller (1938), Clarke stated that this species is “notof frequent occurrence.” NYSM 3731 (Pl. 17, Fig. 5), a frag-ment showing its ribbed inner whorls, might be a Koenen-ites.

Distribution.–Genundewa Limestone (Genesee Group)in Yates and Genesee counties. The lectotype (NYSM 3732)and NYSM 3733 are recorded as from the GenundewaLimestone at Middlesex (Yates County), and NYSM 3731,the possible Koenenites, is from the Portage (Naples) Beds atGriswold (Genesee County), which could be the Genun-dewa Limestone in Murder Creek (Loc. 14a) at Griswold.

Range.–Frasnian: ?Timanites Genozone UD I-?C. ?Re-gional Zone of Manticoceras contratum (17a). ConodontZone uncertain, possibly MN 3.

Manticoceras nodifer (Clarke, 1885)Pl. 18, Figs 11-12; Text-fig. 35B

Goniatites nodifer Clarke, 1885: 21.Manticoceras nodifer. Clarke, 1898, 1899a, b: 74-75, 82, text-figs

54, 63-64, pl. 6, figs 24-26; Miller, 1938: 92-93, pl. 15, figs15-18; House, 1962: text-fig. 3G; Becker & House, 1993:119, pl. 2, figs 2-3.



Type material.–Clarke’s syntypes (USNM 14994a and1499b); lectotype selected by Becker & House (1993).

Diagnosis.–Laterally compressed, subevolute shell withrounded venter, well-rounded umbilical shoulder, and steepumbilical wall. Nodes on inner whorls numbering from 12per volution on inner whorls to 16 on third volution.

Growth lines not observed. Wrinkle layer in lectotypeconsists of fine irregular striae numbering approximately 13per mm on outer flanks at 10 mm diameter, which passbackward from umbilicus and sweep gently forward.

Suture of lecotype (USNM 14994b) at 12 mm diameter,with U-shaped ventrolateral lobe, large asymmetrical lateralsaddle with orad portion of dorsal slope deepest, and broadlyrounded lateral lobe (House, 1962: fig. 3G).

Discussion.–There is some ambiguity as to which speci-men was figured by Clarke (1898, 1899a, b: pl. 6, fig. 26)to show the inner whorls of this species; both specimens arepreserved as internal molds, partly of white calcite, in gray,buff-weathered styliolinid limestone. The drawing appearsto be largely based on the lectotype but it is by no means anaccurate copy because the artist gives the false impressionthat the specimen is perfectly preserved. NYSM 3745 is acast of the lectotype (USNM 14994b) and NYSM 11466(12304/1) is a cast of USNM 14994a.

Distribution.–Upper Genundewa Limestone (GeneseeGroup), Genundewa Point, Canandaigua Lake (YatesCounty; Loc. 23b; Clarke, 1885: 21). The horizon and col-lection date of Clarke’s types are uncertain. The labels withthe specimens indicate their collection by the GeologicalSurvey in 1886 from the “Senecan (Genesee) of LakeCanandaigua.” More recent labels give the horizon as Ge-nundewa. In his original description, Clarke (1885: 21) gavethe explicit location as “Genundewah [original spelling ofGenundewa], Canandaigua Lake.” He described it as a rarespecies and regretted that its “recent discovery” preventedhim from preparing “an illustration of it” in his 1885 report.The horizon within the Genundewa Limestone at Genun-dewa Point (Loc. 23b) is unknown but a position in theupper part of the unit is probable.

Range.–Frasnian: ?Timanites Genozone UD I-?C. ?Re-gional Zone of Manticoceras contractum (17a).

Manticoceras simulator (Hall, 1874)Pl. 9, Figs 4-5

Goniatites simulator Hall, 1974: 2-3; Hall, 1875: 133-134; 1876:pl. 69, figs 1, 3, pl. 74, fig. 8; 1879: 453-455, pl. 69, figs 1-2,pl. 74, fig. 8.

Manticoceras simulator. Hyatt, 1884: 318; House, 1962: text-fig.3F, pl. 45, fig. 14.Type material.–NYSM 3797 (formerly 12309/1), men-

tioned by Hall (1874, 1875) and figured by him (1876: pl.

69, figs 1-2, pl. 74, fig. 8). Stated by Clarke (1899a, b) to bethe only specimen known to Hall; this is the only specimenknown to us.

Dimensions.–Maximum D = 42 mm; D = 37.2 mm;WW = 14.3 mm; WH = 17.2 mm; UW = 7.9 mm.

Diagnosis.–Whorls compressed, very broadly roundedlatterally, narrowly rounded ventrally, deeply impressed dor-sally. Umbilicus deep and moderate in size, less than one-third diameter of shell. Mature suture with rounded laterallobes (Pl. 9, Fig. 5) (House, 1962: fig. 3F). Growth lines notpreserved.

Description.–See Miller (1938: 105) for full description.Discussion.–As noted by Miller (1938), the rounded lat-

eral lobes of the mature suture are distinctive. In typicalspecies assigned toManticoceras, the lateral lobes are narrowto pointed at comparable diameters.

Distribution.–Ithaca Shale and Sandstone (GeneseeGroup), at or near Ithaca (Tompkins County). Precise hori-zon unknown but thought to be from a level above theequivalent of the Lower Genundewa Limestone (Upper Ge-nundewa and West River Shale).

Range.–Frasnian: ?Timanites Genozone UD I-?C. ?Re-gional Zone of Manticoceras contractum (17a) or Koenenitesbeckeri (17b).

Manticoceras sinuosum sinuosum (Hall, 1843)Pl. 9, Figs 6-8, Pl. 10, Figs 1-12, Pl. 11, Figs 1-13, Pl.12, Figs 8-10, Pl. 15, Figs 1-2, Pl. 16, Figs 11-12; Text-

figs 36-38; Table 11.

Goniatites sinuosusHall, 1843: 244, 246, text-figs 106(6), 107(9);1879 (pars): 460-463, pl. 72, fig. 11 (not pl. 70, figs 13, ?14,pl. 74, fig. 11).

Gon. Patersoni (sic) Hall, 1860: 99, text-figs 9-10.Manticoceras PattersoniClarke, 1898, 1899a, b (pars): 45-62, pl. 1,

figs 1-12, pl. 2, figs 1-2, 4 (not figs 3 = M. s. tardum, 6 =Sphaeromanticoceras oxy), pl. 4, figs 16-17, ?18 (not figs 14-15= Probeloceras lutheri?), text-figs 1, 3-11, ?12, 13-28.

Manticoceras sinuosum.Miller, 1938 (pars): 106-115, text-figs 1, 3-11, ?12, 13-28, pl. 18, fig. 4, pl. 19, figs 2-9 (not fig. 1 = M.s. tardum), pl. 20, figs 7-8, ?9, 10 (not fig. 6 = Probeloceraslutheri?); House, 1962: 259.

Type material.–The lectotype of Manticoceras sinuosumsinuosum is Goniatites sinuosus Hall [1843: 243, fig. 106(6)(not 8) = AMNH 5887/1], selected by House (1962: 259),from the Cashaqua Shale, Cashaqua Creek (LivingstonCounty). Among Clarke’s collection of Manticoceras Patter-soni are two fragments from the Naples Beds that have ratherstrong biconvex growth lines with a pronounced lateralsinus, narrow projecting ventrolateral salient and deepventral sinus (hyponomic recurvature): NYSM 3769(Clarke, 1899a, 1899b: pl. 4, fig. 14), Rock Stream (Yates

County), and NYSM 3770 (Clarke, 1899a, 1899b: pl. 4,fig. 15), Naples (Ontario County). This pattern of growthline is not seen in M. s. sinuosum from the Cashaqua at anystage and is more closely comparable with that seen in thelargest specimens of Probeloceras lutheri.

The spelling of pattersoni with two “t”s started withClarke (1898: 59) when he reported Hall as saying it wasintended to honor George Patterson, Lieutenant Gouvernorof the State in 1848, not Paterson’s Gully.

New material.–Most of the specimens reported here arefrom three horizons in the Cashaqua Shale: (1) ParrishLimestone (approximately 60 specimens, mostly fragments);(2) Randall Gully (Ontario County; Loc. 44/4; 10 speci-mens); (3) Shurtleff Septarian Horizon (Livingston and On-tario counties; approximately 50 specimens, all bariticreplacements).

Dimensions.–See Table 11.Diagnosis.–Moderately involute shell with cordate or

heart-shaped cross section. Internal whorls with prominentlamellae (seen in baritic replacements) that become growth-line lirae. Flanks of mature shell sloping gently from maxi-mum whorl width (WW) on the lower flanks to narrowlyrounded venter. Suture with large projecting lateral saddle,acute lateral lobe, and broad umbilical saddle. Biconvex

growth lines weakly sinuous with slight umbilical salient andlateral sinus on mid-flank and distinctive, large, broad ven-trolateral salient.

Description.–Beginning at approximately 5.0 mm diam-eter, whorl compression and impressment of dorsal area in-creasing abruptly; WW/WH ratio rapidly decreasing andUW rapidly increasing (Text-figs 36-38). Typical chordatecross section appearing at approximately 9.0 mm (NYSM13801); WW/WH = 100% by approximately 15 mm di-ameter. WW/WH ratio continuing to decline, but at slowerrate, and leveling off at approximately 35 mm diameter. Atthis stage and beyond, flanks sloping very gradually frommaximum WW on lower flanks to rounded venter (Text-figs 36E, 37A, C). Clarke’s (1898: pl. 4, fig. 16) illustrationof NYSM 3762 (traced as Text-fig. 36A), showing strongconvergence of flanks toward venter, is misleading becauseit is based on a specimen in which the body whorl is moldedin plaster (Pl. 9, Fig. 8). Flanks of mature specimens gentlysloping to narrowly rounded venter (Text-figs 36B-C, G,37A, C, Pl. 10, figs 1, 8.

Suture distinctive, with large, rounded, projecting lateralsaddle, acute lateral lobe, and broad umbilical saddle (Text-figs 37E-H, Pls 9-10).

Elaborate ornamentation lamellae developing after ne-

Table 11. Biometric data for Manticoceras sinuosum sinuosum (Hall, 1843) from the Cashaqua Shale. See Text-fig. 38 for graphs.


NYSM 13835 62.5 20 33.5 9 0.60 0.14NYSM 13800 58.5 ca. 22 ca. 30 ca. 9.5 0.73 0.16

36 14.2 19 6.4 0.75 0.1822 9.5 10.9 4.6 0.87 0.2113.9 6.6 6.5 3.6 1.01 0.268.8 4.4 3.7 2.9 1.19 0.336.0 3.0 2.1 2.3 1.43 0.384.0 2.0 1.4 1.8 1.43 0.452.9 1.4 0.90 1.4 1.56 0.482.1 1.0 0.63 1.0 1.59 0.48

NYSM 13805 54 19 28 12.5 0.68 0.23(CU 42004)NYSM 13809 ca. 48 ca. 18.5 ca. 26.2 8.5 0.71 0.18

ca. 19 8.0 9.5 4.0 0.84 0.21ca. 11.3 ca. 5.5 ca. 5.3 3.1 1.04 0.277.0 ca. 3.5 3.1 2.4 1.13 0.344.6 ca. 2.4 1.7 2.1 1.41 0.463.3 1.5 1.1 1.2 1.36 0.362.4 ca. 1.1 0.75 1.1 1.47 0.46

NYSM 13811 44.4 16 ca. 23 8.0 0.70 0.18NYSM 13801 38 14.7 20 6.4 0.74 0.17

ca. 23 10.2 ca. 11.1 ca. 5.0 0.92 0.2214.6 6.9 7.0 3.8 0.99 0.269.2 4.7 3.9 3.0 1.21 0.336.1 3.3 2.3 2.5 1.43 0.414.2 2.2 1.4 1.9 1.57 0.453.1 1.6 1.0 1.4 1.60 0.452.3 1.1 0.74 1.0 1.49 0.43


pionic constriction (Pl. 11, figs 5-6, 11-13); this feature re-marked upon by Landman et al. (1996: 357-359). Clarke’s(1898, 1899a, b) detailed description of the shell develop-ment and ornamentation of Manticoceras pattersoni was

based on numerous baritic replacements “from scattered cal-careous concretions” in the lower Naples beds between Ho-neoye and Conesus lakes; most probably came from theShurtleff Septarian Horizon of the upper Cashaqua Shale in

Text-fig. 36. Cross sections of Manticoceras sinuosum sinuosum (Hall, 1843) from the Cashaqua Shale. A. NYSM 3762, cross section of aspecimen from the Naples beds, Naples, Ontario County, traced from Clarke (1899a, b: pl. 4, fig. 16; body whorl of specimen modeled inplaster), X 0.7. B-C. NYSM 3773, cross sections of a specimen from the Naples beds, Naples, Ontario County. B. Traced from Clarke(1899a, b: 46, text-fig. 1). C. Traced from a peel of cellophane tape of the same specimen, X 0.8. D-E. NYSM 13801, cross sections of aspecimen from the Parrish Limestone in Conklin Gully (Loc. 47/2). D. (image reversed), X 3.6. E. X 1.4. F-G. NYSM 13800, cross sectionincluding portion of whorl beyond 36.2 mm diameter, from the Parrish Limestone in a tributary to Naples Creek (Loc. 46e/1) above NYRte. 21, near Naples, Ontario County. F. X 3.6. G. X 1.4.


that area. Some of Clarke’s specimens are refigured herealong with newly collected specimens from the ShurtleffSeptarian Horizon. The distinctive ornamentation lamellae,which follow the smooth nepionic stage, gradually evolveinto typical growth lines. Details of their development werecompletely described by Clarke (1898, 1899a, 1889b: 53-54) but it is important to note that the major lamellae arewidely spaced and cross the shell with little or no curvature

to approximately the end of the second whorl; at approxi-mately this stage a shallow sinus (hyponomic sinus) developson the venter and a slight sinus is seen on the mid-flank (Pl.11, Figs 3-8, 11-13). The weakly sinuous, biconvex patternpersists to the largest diameters seen (Pl. 10, Figs 9-11, Pl.11, Figs 1-2). The slight umbilical salient and lateral sinuson the mid-flank and the broad ventrolateral salient are dis-tinctive. Weak revolving lines on the lower flanks are seen at

Text-fig. 37. Sutures and cross sections ofManticoceras sinuosum sinuosum (Hall, 1843) from the Cashaqua Shale. A-B. AMNH 5887/1, crosssection and growth line based on the lectotype designated by House (1962), figured by Hall [1843: text-fig. 106(6)], and recorded as fromCashaqua Creek or Keshequa Creek, near Sonyea, Livingston County, X 0.8. C. NYSM 13816, cross section of a specimen at 40.5 mm di-ameter from Cayuga Creek (Loc. 33/5), Cowesville, Wyoming County, X 1.6. D. NYSM 13802, cross section at 17 mm diameter based ona specimen from the Parrish Limestone, intersection of NY Rte. 21 and Bristol Springs Road (Loc. 46f/1), Naples, Ontario County, X 4.E. AMNH 5889/1:1, suture drawn by MRH in 1959 (a pyritic specimen largely disintegrated by 1998) based on a specimen figured by Hall(1879: pl. 72, fig. 3) asGoniatites patersoniClarke, 1898, and recorded as from Paterson's Creek, Livingston County, X 1.6. F. NYSM 13817,reversed suture at 34 mm diameter of a specimen from Beards Creek (Loc. 38/2), near Pine Tavern Corners, Livingstone County, X 1.2. G.NYSM 13805/CU 42004, reversed suture at 55.5 mm diameter based on a specimen collected by Donald Zenger from the Parrish Limestone,in Conklin Gully (Loc. 47/2) near Naples, Ontario County, X 0.8. H. NYSM 13806, reversed suture at ca. 80 mm diameter based on aspecimen from the Parrish Limestone, Rumpas Hill (Loc. 47a/1) near Naples, Ontario County, X 0.8. Scale = 5 mm (C).


Text-fig.38.Graphsshowingtheontogeny

ofM

antic

oceras

sinuo

sum

sinuo

sum(H

all,1843).SeeTable11

forbiom

etricdata.A

.Based

onmeasurementsofthehypotypesfromthe

CashaquaShale.Sm

allerspecimen

basedon

NYSM

3773,figured

Clarke(1898:41,fig.1).Largerspecimen

basedon

NYSM

4762.B

.Based

onspecimensfrom

theCashaqua

Shale,NYSM

13800-13817.C.B

ased

onspecimensfrom

theShurtleffGullyConcretionHorizon,C

ashaquaShale,NYSM

13818-13835.



large diameters (Pl. 10, Fig. 10). The wrinkle layer (Pl. 15)passes from the inside of the flanks of the body chamber(ventral side) to the dorsal side where it rests on top of thepreceding whorl; these ventral and dorsal wrinkle layers, re-spectively, of House (1971) are clearly confluent.

Discussion.–Clarke (1898, 1899a, b) provided a detaileddescription of Manticoceras pattersoni (=M. sinuosum sinuo-sum) that was summarized with full synonymy by Miller(1938). Clarke treated M. sinuosum as the standard expres-sion of the genus Manticoceras in New York. The otherspecies ofManticoceras in New York were described in termsof “variations from this standard” often in terms of phases inontogeny and geographical (ecological) distribution (Clarke,1898: 79-83, text-fig. 63); some are here assigned to otherspecies and to other genera such as Chutoceras, Koenenites,Sphaeromanticoceras, and Carinoceras. Comparatively littleattention was paid to the details of stratigraphic position ofHall’s and Clarke’s specimens and because of major miscor-relations (corrected following Chadwick, 1935b), the rangesof the species ofManticoceras in New York remained uncer-tain (Clarke, 1898: text-fig. 64).

Detailed descriptions are lacking for many species ofManticoceras in general, which makes comparisons difficult.Among European species regarded (directly or indirectly) asclose to or synonymous with Manticoceras sinuosum are: M.intumescens (Beyrich, 1837), M. cordatum (G. & F. Sand-berger, 1850), M. lamed (G. & F. Sandberger, 1850), andM. buchii (d’Archiac & de Verneuil, 1842). House (1962:259) considered M. sinuosum to be closer to M. cordatumthan toM. intumescens; bothM. sinuosum andM. cordatumhave somewhat flattened flanks that converge (but notsteeply) toward the venter.M. intumescens has a more robust,rounded, subquadrate whorl form (Wedekind, 1913: 52).

North American comparable species are Manticocerasregulare Fenton & Fenton, 1924, M. cordiforme Miller,1938, andM. cf. sinuosumHouse & Pedder, 1963.Mantic-oceras cordiforme (=M. septentrionaleMiller, 1938), from theNorthwest Territories, Canada, andM. sinuosum have a sim-ilar growth-line pattern, butM. cordiforme has a more robustwhorl form (House & Pedder, 1963: 521, text-fig. 7). Theshell development and whorl-form of M. cf. sinuosum fromthe Northwest Territories compares closely withM. sinuosumbut there are differences in details of the suture.

Distribution.–Cashaqua Shale (Sonyea Group), LakeErie (Erie County) to Keuka Lake (Steuben County).Man-ticoceras sinuosum sinuosum occurs in notable abundance inthe Parrish Limestone in the vinicity of Naples in Ontarioand Yates counties (Locs 46, 46a-f, 47, 47a-f ), in concretionhorizons in Barnes Gully, Canandaigua Lake (OntarioCounty; Loc. 44a/2), Randall Gully (Loc. 44/4), south ofBristol Center (Ontario County), and in the Shurtleff Sep-

tarian Horizon, between Conesus and Honeoye lakes (Liv-ingston and Ontario counties; Locs 40-42).

Range.–Frasnian: Probeloceras and ProchoritesGenozonesUD I-E, F. Regional Zones of Probeloceras lutheri (19) andProchorites alveolatus (20). Conodont Zones MN 5-6.

Manticoceras sinuosum clausium n. ssp.Pl. 12, Figs 1-7, 11, Pl. 13, Figs 1-15; Text-figs 39A-I,

40A-B; Table 12

Etymology.–Latin clausium, a more enclosed place, refer-ring to the tighter coiling in the early stages compared withthe nominate subspecies.

Type material.–Holotype NYSM 13839 (Pl. 12, Figs 3-4), Cashaqua Shale, Randall Gully (Ontario County; Loc.44/3).

Material.–Seventy specimens from the lower part of theCashaqua Shale at Randall Gully (Loc. 44/3), 2.5 mi southof Bristol Center (Ontario County; 30 specimens), andWhetstone Brook (Loc. 42/5), 2 mi west of Honeoye (On-tario County; 40 specimens, including baritic replacements).

Diagnosis.–Similar to Manticoceras sinuosum sinuosumbut shell more tightly coiled in early stages, lateral lobe lessacute and lower on flanks, and ornamentation lamellae lessprominent, more closely spaced. and becoming biconvex atearlier stage.

Description.–Tighter coiling (lower UW/D) most appar-ent between 3 and 10 mm diameter (Text-figs 39A-F); shelldevelopment similar to that of Manticoceras sinuosum sinu-osum beyond 15 mm diameter. Lateral lobe (Pl. 12, Fig. 5,Pl. 13, Fig. 12) still rounded and positioned just ventrad ofumbilical shoulder at diameters at which it is acute andhigher on flank inM. sinuosum sinuosum; also less of umbil-ical saddle seen than in M. sinuosum sinuosum.

Juvenile ornamentation lamellae less ornate, becomingsmaller, more sinuous, and closely spaced earlier than innominate subspecies. Lateral and ventral sinuses developingin second volution; growth lines biconvex by start of thirdvolution (Pl. 12, Figs. 7, 11, Pl. 13, Figs. 1, 10). In M. sin-uosum sinuosum, weak biconvex growth lines develop later inthird volution (Pl. 11, Fig. 1).

Discussion.–Manticoceras sinuosum clausium n. ssp. hasonly been found in the lower Cashaqua Shale at two locali-ties in Ontario County. Specimens of Manticoceras fromscattered horizons in the Cashaqua generally show betteragreement with the common nominate subspecies. Varia-tions in tightness of coiling (UW/D) do not seem to followa stratigraphic sequence. For example, specimens of M. s.sinuosum from the Shurtleff Septarian Horizon of the upperCashaqua in Randall Gully (Loc. 44/4) are more similar toM. s. clausium n. ssp. in this feature than specimens from

the intervening Parrish Limestone.Manticoceras sinuosum clausium n. ssp. would seem a log-

ical ancestor ofM. s. sinuosum given its less advanced suture(less acute and more rounded lateral lobe) and earlier strati-graphic appearance in the lower Cashaqua Shale. It is alsotempting to place M. s. clausium n. ssp. in a phylogeneticline between M. simulator from the underlying GeneseeGroup and M. s. sinuosum from the upper Cashaqua Shale.Too little, however, is known about M. simulator; the holo-type (NYSM 3797) and only known specimen is from anunknown horizon and locality (probably in the upper Gene-see Group) near Ithaca (Tompkins County). Manticocerassimulator still has a rounded lateral lobe on the umbilicalshoulder at 36.5 mm diameter but the umbilical width atthis diameter is even larger than in M. s. sinuosum.

Manticoceras sinuosum clausium n. ssp. could be moreclosely related toM. contractum from the Genundewa Lime-stone (middle Genesee Group); unfortunately two of thethree figured syntypes are small, and nothing is known ofthe suture. The umbilicus of M. contractum is narrow andthe whorl form similar (Clarke, 1898, 1899a, b: text-fig. 44)but slightly more compressed; its growth lines are distinctlybiconvex by the end of the second volution (Miller, 1938:82) as inM. s. clausium n. ssp., but the lamellae tend to clus-ter in pairs, forming fascicles.

Distribution.–Lower Cashaqua Shale (Sonyea Group) atWhetstone Brook, (Loc. 42/5) near Honeoye (OntarioCounty), and Randall Gully (Loc. 44/3) near Bristol Center(Ontario County).

Range.–Frasnian: Probeloceras Genozone UD I-E. Re-gional Zone of P. lutheri (19). Conodont Zone MN 5.

Manticoceras sinuosum tardum Clarke, 1898Pl. 9, Figs 2-3, Pl. 14, Figs 1-10; Text-figs 39J, 40C;

Table 13

Manticoceras tardum Clarke, 1898, 1899a, b: 63-64, 81, pl. 1, fig.13.

?Manticoceras tardumClarke, 1898, 1899a, b: pl. 6, fig. 31 (NYSM3804).

Manticoceras PattersoniClarke, 1898, 1899a, b (pars): 45-62, pl. 2,fig. 3.

Manticoceras tardum. Miller, 1938 (pars): 121-123, pl. 24, fig. 6.(?) Manticoceras tardum. Miller, 1938: pl. 24, fig. 7.Manticoceras sinuosumMiller, 1938 (pars): 106-115, pl. 19, fig. 1.

Type material.—NYSM 3804, a plastotype, the largerand surviving syntype figured by Clarke, is here designatedas lectotype. A specimen figured as Manticoceras Pattersoniby Clarke (1898, 1899a, b: pl. 2, fig. 3; NYSM 3766) is in-cluded in the subspecies.

New material.–Ten specimens (mostly small baritic re-placements) from the Cashaqua Shale, Briggs Gully (Loc.43/3), Honeoye Lake. Bed 43/3 is believed to be the sourceof the smaller syntype figured by Clarke (1898, 1899a, b: pl.1, fig. 13), which is lost (Miller, 1938: 123).

Dimensions.–See Table 13.Discussion.–Clarke (1898, 1899a, b) describedMantico-

ceras tardum as a species with a “primitive” ornamentation in

Table 12. Biometric data for Manticoceras sinuosum clausium n. ssp. from the Cashaqua Shale. See Text-fig. 40 for graphs.


NYSM 13839 41 15.7 22 7.0 0.71 0.17(holotype)

NYSM 13857 40.4 15.5 ca. 20.4 ca. 7.4 0.76 0.18NYSM 13848 27.5 11.0 14.0 5.0 0.79 0.18

17.0 8.0 8.7 3.7 0.92 0.22ca. 10.3 5.2 4.7 3.0 1.11 0.296.6 3.6 2.9 2.1 1.24 0.324.3 2.4 1.7 1.5 1.41 0.353.0 1.5 1.0 1.3 1.50 0.432.1 1.0 0.67 — 1.49 —

NYSM 13842 23.0 ca. 10.0 11.5 ca. 4.0 0.87 0.17NYSM 13849 16.6 7.7 8.3 3.0 0.93 0.18

10.2 5.4 5.1 2.2 1.06 0.226.2 3.5 2.8 1.7 1.25 0.273.9 2.3 1.7 1.4 1.35 0.362.6 1.3 0.90 1.0 1.44 0.38

NYSM 13854 10.3 5.7 5.0 ca. 2.3 1.14 0.226.4 3.8 2.9 1.9 1.31 0.304.0 2.4 1.7 1.5 1.41 0.382.7 1.5 0.96 1.1 1.56 0.411.9 1.0 0.64 0.80 1.56 0.42


which simple distant ornamentation lamellae persist withoutintercalation to the end of the third whorl and a much widerumbilicus and also broader venter than M. pattersoni (= M.sinuosum). The locality given for the syntypes was the lowerPortage shales of Briggs Gully; Miller (1938: 123) stated thatthey came only from the Cashaqua Shale at that locality.The smaller of the syntypes (now lost) was a baritic replace-

ment of approximately three whorls; the larger of the syn-types (NYSM 3804) is a plastotype of four whorls.

The protoconch is similar to that of Manticoceras sinuo-sum, but appears to be more globular in shape, as the whorlsection is strongly reduced after the nepionic swelling (Pl.14,figs 1, 3). The first few whorls are rather flattened dorsovent-rally, producing a broad venter, and narrowly rounded flanks

Text-fig. 39. Sutures and cross sections of Manticoceras sinuosum clausium n. ssp. and M. s. tardum Clarke, 1898, from the lower CashaquaShale. A-I. M. s. clausium n. ssp., from Randall Gully (Loc. 44/3) near Bristol Center, Ontario County, unless otherwise stated. A. NYSM13849, cross section, including portion of whorl beyond, at 16.6 mm diameter, X 1.5. B. NYSM 13851, cross section, X 1.5. C. NYSM13852, cross section at 5.7 mm diameter, X 3.7. D. NYSM 13813, cross section, based on a specimen from Eighteenmile Creek (Loc.30/10), North Evans, Erie County, X 3.7. E-F. NYSM 13847, cross sections, X 1.5 (E), X 3.7 (F). G. NYSM 13846, reversed suture at 17.5mm diameter, X 3.7. H. NYSM 13842, suture at 23 mm diameter, X 3.7. I. NYSM 13839, suture at ca. 30 mm diameter based on the holo-type, X 1.1. J. M. s. tardum, NYSM 13870, cross section at 82 mm diameter based on a specimen from Briggs Gully (Loc. 43/3), HoneoyeLake, Ontario County, X 0.9.


and slight dorsal impressment.Simple distinct growth lamellae are still seen in NYSM

13879 (Pl. 14, Fig. 1) at the end of three whorls. Low trans-verse ribs, which cross the flanks and venter, are seen in in-ternal molds (Pl. 14, Figs. 2, 4, 6-7); the ribs and interveningbroad depressions are traversed by fine, closely spaced stria-tions. The major growth lamellae of the shell wall (Pl. 14,Figs 4, 6-7) overlie the ribs and depressions.

The umbilical widths of the smaller specimens are onlyslightly larger than in Manticoceras sinuosum sinuosum, andthey compare with the smaller of Clarke’s syntypes. Thewhorl form and umbilical widths of the larger specimens,however, cannot be distinguished from the nominate sub-species at the same stage (Text-fig. 39J).

The suture is seen near the end of the second whorl, andhere the ventral lobe is trifid, as in Manticoceras sinuosumsinuosum; the suture and growth lines of NYSM 13843 at 17mm diameter also cannot be distinguished from the nomi-nate subspecies at the same stage.

The most serious problem in determining Manticocerassinuosum tardum concerns the degree of umbilication in thetwo syntypes. Clarke (1898, 1899a, b: 63) noted: “The um-bilication ofM. tardum is much greater than that ofM. Pat-tersoni and M. apprimatum. . . .The form and crosssectionof the whorl is distinct from both at parallel growth stages,being broader and flatter on the venter.” Miller (1938: 122)examined the larger syntype (NYSM 3804) and cited thefollowing dimensions: D = ca. 11.5 mm; UW = 6.5 mm;WH = 4 mm. The UW value is significantly greater than inM. sinuosum sinosum and the specimens in hand. The UWof the smaller syntype figured by Clarke, however, compareswith specimens of M. pattersoni of similar size on the sameplate.

It is possible that NYSM 3804 came from a differenthorizon and is not the same species as the smaller figured

specimen. This is apparently the case with Manticoceras ap-primatum. Here the wide umbilicus and simple persistantgrowth lamellae of the smallest syntype (NYSM 3731; alsoa plastotype) compare closely with M. sinuosum tardum(NYSM 3804) figured on the same plate. NYSM 3731 isfrom the Naples beds (Cashaqua?), at Griswold. Based onthe two larger “syntypes” from the Genundewa Limestone,however,M. apprimatum is distinguished by its very narrowumbilicus: “The umbilicus is very deep and narrow, closerthan in any other species, even more extreme than in Man-ticoceras contractum.” (Clarke, 1898, 1899a, b: 62).

The new specimens are conspecific with the smaller ofthe figured specimens of Manticoceras sinuosum tardum,based on their similar growth-line lamellae, degree of umbil-ication, and rather broad, flattened venter in the earlywhorls. The presence of larger forms that are indistinguish-able fromM. s. sinuosum suggests thatM. tardum is best re-garded as a subspecies of M. sinuosum.

Distribution.–Lower Cashaqua Shale (Sonyea Group) atBriggs Gully, Honeoye Lake (Ontario County; Loc. 43/3).

Range.–Frasnian: Probeloceras Genozone UD I-E. Re-gional Zone of P. lutheri (19). Conodont Zone MN 5.

Manticoceras lamed (G. & F. Sandberger, 1850)Pl. 16, Figs 1-2, 9-10; Text-figs 41A-C, 42E; Table 14

Goniatites lamed var. complanatusG. & F. Sandberger 1850: 90, pl.8, figs 5, 5a; Wedekind, 1913: pl. 6, fig. 3.

Manticoceras lamedMiller, 1932: 331; House & Ziegler 1977: 83,pl. 2, figs 15-17, 23-24.

Manticoceras (Manticoceras) cordatum (pars) Clausen, 1969: 130.

Type Material.–A lectotype was chosen and illustrated byHouse [in House & Ziegler, 1977: pl. 2, figs 15-17, BerlinMuseum (HUM) c. 491] from among the specimens illus-

Table 13. Biometric data for Manticoceras sinuosum tardum Clarke, 1898, from the Cashaqua Shale. See Text-fig. 40 for graph.


NYSM 13870 ca. 82 28.0 41.0 14.3 0.68 0.17ca. 53.5 — — 9.0 — 0.17ca. 33 ca. 13.0 17.0 ca. 6.6 0.76 0.20ca. 20.5 — — ca. 5.5 — 0.27

NYSM 13873 17.0 7.1 7.0 ca. 5.0 1.01 0.29NYSM 13872 ca. 5.6 — — 2.4 — 0.43

4.1 2.0 1.3 1.9 1.54 0.46ca. 2.9 — — 1.4 — 0.482.2 0.96 0.65 1.0 1.48 0.45

NYSM 13871 5.4 2.8 2.0 2.3 1.40 0.433.8 1.8 1.2 ca. 1.7 1.50 0.452.8 1.3 ca. 0.83 ca. 1.3 1.57 0.462.0 0.90 ca. 0.62 0.94 1.45 0.74

NYSM 13874 5.0 2.2 ca. 1.7 ca. 2.4 1.29 0.48


Text-fig.40.Graphsshow

ingtheontogeny

ofthesubspeciesof

Man

ticoceras

sinuo

sum(H

all,1843).SeeTables12

and13

forbiom

etricdata.A

-B.M

.s.c

lausiu

mn.ssp.,based

onmeasurementsofspecimensfrom

theCashaquaShale.A.N

YSM

13839and13848-13849.B.13840-13847,13854-13868.C

.M.s

.tar

dum,based

onmeasurementsofNYSM

13869-13874.


trated by the Sandberger brothers; another of their speci-mens was also figured (pl. 2, figs 23-24). The type materialwas from several German localities and the source of the lec-totype is not clear.

New material.–Seventeen specimens (NYSM 12057-12059, 12062-12067, and others) preserved in mudrockand crystalline calcite mostly as internal molds but in somewith the shell as a crystalline calcite replacement. Phragmo-cones and incomplete body chambers are represented.

Dimensions.–The dimensions of the lectotype [BerlinMuseum (HUM) c. 491] are as follows: D = 22.4 mm,WW= 9.3 mm;WH = 12.0 mm; UW = ca. 3.5 mm. For dimen-sions of the New York material, see Table 14.

Description.–Openly umbilicate in early whorls (Text-fig. 42E) but in middle and outer whorls subinvolute withrelatively small umbilicus. Maximum whorl width close torounded umbilical shoulder; flanks converging in smoothcurve to well-rounded to narrowly rounded venter. Suturesas illustrated in Text-fig. 41, often showing no subdivisionof mid-ventral saddle in early whorls, but later saddle be-coming divided and increasing markedly in prominence rel-ative to lateral saddle, which shows ratio change from ca.54% at 19 mm diameter to ca. 98% at approximately 68mm diameter (Text-figs 41A-B). Growth lines not seen inlarge specimens, but in middle whorls show broad, flat lat-eral sinus passing with slight convexity to ventrolateralsalient and back to well-rounded ventral sinus.

Discussion.–The nameManticoceras lamed is given prece-dence over M. cordatum and the latter considered as a sub-species of M. lamed (see House & Ziegler, 1977). Alectotype for M. cordatum was designated by House (inHouse & Ziegler, 1977: pl. 2, figs 13-14), which is now thelectotype of the subspecies, a stouter variety. One New Yorkspecimen (NYSM 12057; Pl. 16, Figs 9-10) could be refer-able to the stouter subspecies.

These rather laterally compressed New York manticocer-atids with an elegantly ovoid cross section are close toMan-ticoceras sinuosum, and much material could beindistinguishable from that species. The lectotype of M. s.

Table 14. Biometric data for Manticoceras lamed (G. & F. Sandberger, 1850) from the Rhinestreet and Angola shales.


NYSM 12057 94.3 33.2 48.9 ca. 17.6 0.68 0.19NYSM 12058 23.7 9.2 — — — —NYSM 12067 16.4 7.3 — — — —NYSM 12059 16.2 7.5 — — — —NYSM 12066 16.2 ca. 7.2 8.3 3.6 0.87 0.22NYSM 12065 10.9 5.1 5.6 2.5 0.91 0.23NYSM 12064 10.0 5.0 ca. 4.8 — 1.04 —NYSM 12063 7.5 3.8 ca. 3.8 ca. 1.2 1.00 0.16NYSM 12062 5.6 3.3 ca. 3.4 — 0.97 —

Text-fig. 41. Sutures ofManticoceras, Sphaeromanticoceras, andDel-phiceras from the West Falls Group in New York State. A-C. M.lamed (G. & F. Sandberger, 1850). A. NYSM 12057, suture at anestimated 68 mm diameter based on a specimen from AngolaShale equivalents in Wolf Creek (Loc. 85), Castile, WyomingCounty, X 0.7. B. NYSM 12058, suture at ca. 19 mm diameterbased on a specimen from the upper Rhinestreet Shale in JohnsonCreek (Loc. 58/1),Wyoming County, X 1.9. C. NYSM 12059, su-ture at ca. 6.5 mm diameter based on a specimen from lower An-gola Shale equivalents at Varysburg (Loc. 79/4),Wyoming County,X 3.1. D-E.M. aff. lamed, NYSM 12060. D. Suture at ca. 11 mmdiameter, based on a specimen from the Point Breeze GoniatiteBed, lower Angola Shale, in Relyea Creek (Loc. 82/6), WyomingCounty, X 1.9. E. Growth line at ca. 17 mm diameter, based onsame specimen as D. F. S. rhynchostomum (Clarke, 1898), NYSM12061, suture based on an incomplete specimen from equivalentsof the upper Rhinestreet Shale, in the probable Relyea Creek Hori-zon, in Kennedy Gulf (Loc. 63/3), near Dale, Wyoming County,X 0.7. G.D. cataphractum (Clarke, 1898), NYSM 3643, lectotypehere designated, suture at 8 mm diameter, of a specimen recordedas from the Hanover Shale, Java, Wyoming County, X 6.4.


sinuosum, however, shows a growth line with scarcely a lat-eral sinus at all; as described previously, early stages of Hall’sspecies seem to divide the ventral median saddle earlier.

This is a common Frasnian species in Europe, but unfor-tunately none of the European successions have provided asatisfactory time sequence of material. The Büdesheim spec-imens described by Clausen (1969), are mostly very small,so that it is difficult to compare with the Manticoceras sinu-osum and M. aff. lamed material described here from NewYork. Manticoceras lamed is described from Adorf, RhenishSchifergebirge (House & Ziegler, 1977: 83) only from thenodulosum Zone (lower cordatum Zone), and there was apreference for using a German species name rather than im-porting the American name M. sinuosum for that material.

More work is needed on the European material before allthis can be resolved.

Distribution.–Upper Rhinestreet Shale and Angola Shale(West Falls Group) and equivalents in Erie and Wyomingcounties. Specimens from the upper Rhinestreet Shale in-clude: Cazenovia Creek (Erie County; Loc. 54/1), 3196/23;Varysburg (Wyoming County; Loc. 57/2), NYSM 12064,12065, and 12067; Johnson Creek (Wyoming County; Loc.58/1), NYSM 12058; Relyea Creek (Wyoming County;Loc. 60/4), 3193/5; Kennedy Gulf (Wyoming County; Loc.63/3), 3234/2.

Specimens from the lower Angola Shale include: Hamp-ton Brook (Erie County; Loc. 74/4), 3274/4; CazenoviaCreek (Erie County; Loc. 75/4), 3251/1a; Sheldon Creek

Text-fig. 42. Cross sections of Playfordites, Sphaeromanticoceras, Carinoceras, and Manticoceras from New York State. A. P. cf. tripartitus (G.& F. Sandberger, 1850), NYSM 12074, cross section based on a specimen from equivalents of the upper Rhinestreet Shale in Johnson Creek(Loc. 58/1), Wyoming County, X 4.3. B. S. rhynchostomum (Clarke, 1898), NYSM 12085, from equivalents of the upper Rhinestreet Shalein Johnson Creek (Loc. 58/1), Wyoming County, X 2. C. C. vagans (Clarke, 1898), NYSM 12072, from the lower Angola Shale in HamptonBrook (Loc. 74/3), Erie County, X 2. D. S. oxy (Clarke, 1897), NYSM 12036, from the Point Breeze Goniatite Bed, lower Angola Shale inBig Sister Creek (Loc. 73/6), Angola, Erie County, X 2. E. M. lamed (G. & F. Sandberger, 1850), NYSM 12058, from upper RhinestreetShale equivalents in Johnson Creek (Loc. 58/1), Wyoming County, X 1.9. F. M. sinuosum apprimatum (Clarke, 1898), NYSM 3732, thelectotype, from the Genundewa Limestone, Middlesex, Yates County, X 2. G. C. sororium (Clarke, 1898), NYSM 3803, new cross sectionthought to be a cotype figured by Clarke (1899a, b: 76, fig. 59), believed to be former NYSM 12310/6, probably from the lower AngolaShale, near Angola, Erie County, X 2. H-I. S. rhynchostomum. H. NYSM 12073, from the Point Breeze Goniatite Bed of the lower AngolaShale in Relyea Creek (Loc. 82/6), Wyoming County, X 2. I. NYSM 12088a, from the lower Angola Shale in Big Sister Creek (Loc. 73/6),Angola, Erie County, X 2.


(Wyoming County; Loc. 78/4b), 3226/9; Varysburg(Wyoming County; Loc. 79/4), NYSM 12059, 12063,3214/4.

Specimens from the upper Angola Shale include: LakeErie Shore (Farnham Creek) (Loc. 71/9a), NYSM 12066;Big Sister Creek (Erie County; Loc. 73/9), NYSM 12062;Varysburg (Wyoming County; Loc. 79/10), 3215/12; WolfCreek (Wyoming County; Loc. 85), NYSM 12057.

Range.–Frasnian: ?Late Beloceras-Playfordites-Neoman-ticocerasGenozones UD-I, ?H-J. Regional Zone of ?Schinde-wolfoceras chemungensis to Sphaeromanticocerasrhynchostomum (?21c-22b). Conodont zones very uncertain,?MN 8-11.

Manticoceras lamed aff. cordatum(G. & F. Sandberger, 1850)

Text-figs 33K-L

Material.–Three fragments (NYSM 16568, 3911/14,3911/18) of calcitic replacements from the Fossil Log Hori-zon of the upper West River Shale (the Williamsburgh Bedof Over et al., 1999, 2003), 3.4 m below the MiddlesexShale in Beards Creek, Leicester (Livingston County; Bed18a/4b).

Discussion.–Assignment to the subspecies Manticocerascordatum lineage (House & Ziegler, 1977) is based primarilyon whorl form. The shell is widest across the lower flanks,and the flanks slope gently to a broad, rounded venter (in-cluding NYSM 16568 at ca. 9.3 mm diameter). In cross sec-tion, they resemble the rather tubby M. contractum of theGenundewa Limestone, but because the growth lines are notpreserved, comparison with the distinctive pattern in thatspecies cannot be made.

The suture of NYSM 16568, one-half whorl back from12.5 mm diameter, shows a large, broad lateral saddle and abroadly rounded lateral lobe.

Distribution.–UpperWest River Shale (Genesee Group),Fossil Log Horizon (Williamsburgh Bed of Over et al.,1999, 2003), Beards Creek, Leicester (Livingston County;Bed 18a/4b).

Range.–Frasnian: Timanites Genozone UD I-C. Re-gional Zone of Koenenites beckeri (17b). Conodont Zone

MN 4.

Manticoceras aff. lamed (G. & F. Sandberger, 1850)Pl. 16, Figs 3-8; Text-figs 41D-E; Table 15

aff. Goniatites lamed var. complanatus G. & F. Sandberger 1850:90, pl. 8, figs 5, 5a.

Material.–Seven specimens referred to below and twoothers all preserved in mudrock and crystalline calcite assolid specimens often slightly crushed in the outer whorlswith shell preserved where present as recrystallized calcite, allfrom the Point Breeze Goniatite Bed in the lower AngolaShale.

Dimensions.–See Table 15.Description.–Subinvolute with maximum whorl width

close to rounded umbilical shoulder with flanks convergingflatly to ventrolateral area with slight concavity in outer partoften emphasized by diagenetic crushing; rounded butabrupt turnover of flanks to flatly rounded, wide venter. Su-ture (Text-fig. 41D) showing distinctive, low, divided ventralsaddle, broad lateral saddle, and wide, rounded lateral lobeat 11 mm diameter. Growth lines weakly biconvex (Text-fig. 41E; Pl. 16, Fig. 7) with broad, flat, slightly prosiradiatesinus on flanks, asymmetric ventrolateral salient, and broad,rounded sinus on venter. Growth-line striae prominent (Pl.16, Figs 3, 5, 7) and often bunched, simulating weak, fineribbing on outer whorls (NYSM 12060 and 12068).

Discussion.–As noted above, Manticoceras lamed isrecorded from the upper Rhinestreet Shale to the upper An-gola Shale, but not from the Point Breeze Goniatite Bed,which yields the richest fauna of the lower Angola Shale.The form here described as M. aff. lamed is obviously partof the M. lamed lineage, but the differences are sufficientlyclear cut to draw attention to them by the use of opennomenclature because there is a greater stoutness to thewhorl section and a more abrupt turnover from the flanks toa wider rounded venter than in M. lamed, from which theyalso differ in lacking the elegant whorl outline of the longerranging form. The whorl form approaches that ofM. schell-wieniWedekind (1913: 65, pl. 5, figs 1-2) but the holotypeof that species is noticeably more compressed (at D = 27

Table 15. Biometric data for Manticoceras aff. lamed (G. & F. Sandberger, 1850) from the Angola Shale.


NYSM 12068 22.9 8.7 11.5 3.1 0.76 0.14NYSM 12069 ca. 18.1 7.6 9.5 3.0 0.80 0.17NYSM 12060 16.8 7.7 7.2 ca. 2.5 1.07 0.15NYSM 12070 12.6 5.2 ca. 6.6 ca. 2.5 0.79 0.20NYSM 12071 12.4 5.9 7.1 — 0.83 —



mm, WW = 7.6 mm) and the flanks are rather flatter andmore parallel sided than the New York material.

Distribution.–Known only from the Point Breeze Goni-atite Bed in the lower Angola Shale (West Falls Group) inWyoming County, at Relyea Creek (Loc. 82/6), NYSM12060, 12068-12070, 3211/1a, and 3212/7, and StonyCreek (Loc. 83/6), NYSM 12071.

Range.–Frasnian: Neomanticoceras Genozone UD I-J.Regional Zone of Sphaeromanticoceras rhynchostomum (22b).Conodont Zone ?MN 11.

Manticoceras sp.Pl. 17, Fig. 4

Manticoceras acceleransClarke, 1897: 53, nom. nud.; Clarke, 1898:53; 1899a, b: 77, 82, text-figs 60, 63-64, pl. 6, fig. 10; Miller,1938: 74, text-fig. 12, pl. 13, fig. 8.

Type Material.–The only available syntype of Mantico-ceras accelerans, here designated lectotype, is NYSM 3730(formerly 12300/1).

Description.–See accounts of Clarke (1898, 1899a, b: 77)and Miller, (1938: 74).

Discussion.–We regard this crushed and unsatisfactorylectotype as unusable for taxonomic purposes. However thegeneric assignment seems probable.

Distribution.–Reported to be from Cashaqua Shale(Sonyea Group) at Naples (Ontario County).

Range.–Frasnian: Probeloceras or Prochorites GenozoneUD I-E or F. Regional Zone of Probeloceras lutheri (19) orProchorites alveolatus (20). Conodont Zone MN 5 or 6.

Genus CARINOCERAS Ljaschenko, 1957Type species.–Carinoceras menneri Ljaschenko (1957:

199), by original designation.Diagnosis.–Gephuroceratids with oxyconic, compressed,

and often keeled outer whorls; innermost whorls evolute;middle whorls oxyconic to sagittate or rounded, subinvo-lute. Reaching oxyconic form at early stage. Adult with deep,narrow umbilicus. Suture as in Manticoceras or, as in typespecies, with incipient subdivision of ventral lobe in outerwhorls. Shell surface smooth, growth lines biconvex or con-vex.

Discussion.–Genus comprises two groups of species. Thefirst with biconvex growth lines includes: Carinoceras men-neri Ljashenko, 1957: 199 (pars), pl. 1, fig. 2; C. galeatum(Wedekind, 1913: 60, pl. 4, figs 3-4, text-fig. 8a); C. soror-ium (Clarke, 1898, 1899a, b: 75, 82, pl. 4, figs 1-5, text-figs 56-59, 63-64; cotypes NYSM 3800-3802); and C.vagans (Clarke, 1898, 1899a, b: 78, 82, pl. 6, figs 11-12,text-figs 62-64; holotype NYSM 3805).

The second group appears to have convex growth lines

and includes: C. acutum (G. & F. Sandberger, 1850: pl. 7,figs 1, 1a, 1b; 1851: 82); C. acutiforme (H. & G. Termier,1950: 53, 151, text-figs 13-14). Note that Manticoceras oxyis here referred to Sphaeromanticoceras because it is shownthat the discoidal form is a late stage development.

Oxyconic form is an iteratively repeated ammonoidmorphology (House, 1981a: 17) and there is no reason tosuppose it arose only once from Manticoceras in the Ge-phuroceratidae. Nevertheless, these two groups of speciesform a compact group. Possible derivations of it are formssuch as M. inversum (Wedekind, 1913) and M. carinatum(Beyrich, 1837), which are evolute in innermost whorls,oxyconic or carinate and sagittate in their middle whorls,but that revert to a typical M. lamed-cordatum type withrounded outline in outer whorls.

Separation into two groups here is to emphasize the twodifferent patterns of growth-line form in mature individuals.In the second group, there is only a lateral salient, more lat-eral than ventrolateral in position. In the first group, thereare generally biconvex growth lines, but the original figuresof Costamanticoceras koeneni (Holzapfel, 1882) show bicon-vex growth lines in early whorls and then an increasingprominence of a broad ventrolateral salient (Holzapfel,1882: pl. 46, figs 4, 4b, 6) that suggests an intermediate po-sition (see Becker & House, 1993), and even the holotypeof C. vagans described here shows a similar tendancy to abroad and prominent ventrolateral salient although thegrowth lines as a whole are biconvex; both of these speciesare substantially smaller than specimens referred to the sec-ond group.

Distribution.–North America, Europe, Russia, NorthAfrica, and Australia.

Range.–Frasnian: Prochorites Genozone to Neomantico-ceras Genozone UD I-F to J; possibly to Archoceras andCrickites Genozones (UD I-K to L).

Carinoceras sororium (Clarke, 1898)Pl. 18, Figs 3-4, 6-9; Text-fig. 42G

Manticoceras sororium Clarke, 1898, 1899a, b: 75-76, 82, text-figs56-59, pl. 4, figs 1-5; Frech, 1913: 24; Miller, 1938: 117-119,text-figs 23A-D, pl. 24, figs 1-5.

Manticoceras (Carinoceras) sororium. Clausen, 1971: 182, 198.Carinoceras sororium. House & Kirchgasser, 1993: 276.

Type material.–Lectotype here selected as NYSM 3802(Pl. 18, Figs 8-9). Original specimens (NYSM 3798-3803)are gutta percha molds obtained after dissolution of calciticreplacements. This method of extraction shows the shell or-nament superbly, but destroys other shell characteristics.

Dimensions.–The dimensions of figured material ofClarke (1899a, b) are as follows: NYSM 3798 (pl. 4, fig. 1),

D = 6 mm; NYSM 3799 (pl. 4, figs 2-3), D = 5.4 mm;NYSM 3800 (pl. 4, fig. 4), D = 11.2 mm; NYSM 3801 (pl.4, fig. 5), D = 11.3 mm.

Description.–Detailed descriptions were given by Clarke(1899a, b: 75-76, 82) andMiller (1938: 117-119) and neednot be repeated. The original material is partly reillustratedphotographically here for the first time (Pl. 18, Figs 3-4, 6-9). The technique of mold preparation leads to uncertaintyon the precise shape of outer whorls.

Discussion.—This species has not been identified in ourcollections. Clarke, however, illustrated distinctive earlystages. It is considered that it might represent early stagessimilar to Carinoceras vagans but an adult with roundedrather than acute venters.

Distribution.–NYSM 3798-3803 are recorded from “thecalcareous concretionary masses in the shales on Big SisterCreek, Erie County” (Clarke, 1899c: 106-107) and all spec-imens known to Clarke were from the vicinity of Angola(Loc. 73). The source of the material is probably all fromwithin the lower Angola Shale. Clarke mentioned FarnhamCreek as another locality (Locs 71-72).

Distribution.–Lower Angola Shale (West Falls Group;Erie County).

Range.–Frasnian: Neomanticoceras Genozone UD I-J.

Carinoceras vagans (Clarke, 1898)Pl. 23, Figs 5-16; Text-figs 43A, D-E, 44D, 46A; Table 16

Manticoceras vagans Clarke, 1898, 1899a, b: 78-79, 82, text-fig.62, pl. 6, figs 11-12; Frech, 1913: 24; Miller, 1938: pl. 15, figs11-14; House, 1962: 258-259.

Manticoceras (Carinoceras) vagans. Clausen 1971: 182, 198.Carinoceras vagans. House & Kirchgasser, 1993: 276.Carinoceras oxy House & Kirchgasser, 1993: figs 4E-F.

Type Material.–Only one specimen, the holotype(NYSM 3805), was known to Clarke (1898,1899a, b) andto Miller (1938), who each described it. It is illustrated pho-

tographically for the first time here (Pl. 23, Figs 13-14) anda new preparation has been made of the suture (Text-fig.43A).

New Material.–Eleven specimens (including NYSM12036-12042 and 12048) have been found in situ duringthis work, all from the upper Rhinestreet Shale and AngolaShale. The specimens are mostly small but preserved solid inmudstone often with growth lines and frequently with su-tures shown.

Dimensions.–See Table 16 and Text-fig. 46A.Description.–Innermost couple of volutions evolute;

whorl section rounded, depressed; by 6 mm diameter(NYSM 12040) carinate form of adult already seen (Text-fig. 43E) andWH =WW at approximately 12 mm diameter(NYSM 12040); thereafter whorl height increasing relativelyrapidly (Text-fig. 46A). Maximum whorl width at 18 mmdiameter close to umbilical shoulder and flanks convergeflatly, but with slight convexity, to acute venter. At approx-imately 20 mm diameter, whorls have maximum width closeto umbilicus and flanks slope convexly and evenly to suba-cute venter. By 27 mm diameter (NYSM 12036), trend con-tinuing with venter so sharply acute to feel knife-edged;suture near this diameter illustrated inText-fig. 44D. Largerspecimens (NYSM 12044) much stouter in cross section butstill galeate. Sutures of earliest whorls unknown. Growthlines biconvex at all stages seen, with shallow latero-umbil-ical salient essentially on shoulder and increasingly broadand prominent ventrolateral salient (Text-fig. 43D); growthlines not seen above 23 mm diameter.

Discussion.–The stratigraphically youngest specimensknown, from above the level of the Point Breeze GoniatiteBed (NYSM 12043-12045), are determined as Carinocerasaff. vagans on account of the rather more robust whorl sec-tion and the more forward position of the lateral lobe.

The group of “species” Carinoceras menneri, C. galeatum,C. vagans, C. acutum, and C. acutiforme share much in com-mon and there is a case to be made for preserving the names

Table 16. Biometric data for Carinoceras vagans (Clarke, 1898) Group from the Rhinestreet and Angola Shales. See Text-fig. 46A forgraph.


NYSM 3805 22.3 10.0 11.6 ca. 4.5 0.86 0.20(holotype)NYSM 12045 39.4 ca. 15.0 21.0 7.4 0.71 0.19NYSM 12044 31.5 ca. 12.3 — — — —NYSM 12036 29.9 8.2 18.2 ca. 3.0 0.45 9.9NYSM 12043 24.6 ca. 11.0 12.0 5.8 0.92 0.24NYSM 12042 15.4 6.5 7.8 -- 0.83 —NYSM 12041 12.8 5.4 6.7 ca. 3.1 0.81 0.24NYSM 12040 11.8 5.8 5.8 ca. 3.2 1.00 0.27NYSM 12039 10.2 4.8 — — — —NYSM 12038 9.6 4.5 ca. 5.0 — 0.90 —NYSM 12037 8.3 3.1 ca. 3.4 2.5 0.91 0.30


Text-fig. 43. Sutures, a growth line, and cross section of Carinoceras vagans (Clarke, 1898) Group and Sphaeromanticoceras rhynchostomum(Clarke, 1898) from the West Falls Group, New York. A. C. vagans, NYSM 3805, suture at ca. 22 mm diameter of the holotype recordedas from "a loose block of sandstone among the Portage outcrops in the town of Naples," X 4.5. B-C. C. aff. vagans. B. NYSM 12045, sutureat ca. 27 mm diameter based on a specimen from Angola Shale equivalents at Varysburg (Loc. 79/10), Wyoming County, X 4.5. C. NYSM12044, suture at ca. 21 mm diameter based on a specimen from above the Point Breeze Goniatite Bed in Angola Shale equivalents at GladeCreek (Loc. 77/1), Wyoming County, the highest specimen of the Group known in New York, X 4.5. D-E. C. vagans. D. NYSM 12048,growth line at 17.3 mm diameter based on a specimen from upper Rhinestreet Shale equivalents on Relyea Creek (Loc. 60/4), WyomingCounty, X 4.5. E. NYSM 12040, cross section of a juvenile at 13.9 mm diameter reconstructed from a specimen from the upper RhinestreetShale equivalents on Johnson Creek (Loc. 58/1), Wyoming County, X 4.1. F-G. S. rhynchostomum,NYSM 12035, suture at WH = 64 mmwith very high median saddle and growth lines at WH = 61 mm, based on a specimen from the Angola Shale on Hampton Brook (Loc.74/6a), Erie County, X 1.8.


merely as subspecies within the oldest available name, C.acutum. Comparisons are aggravated by the lack of muchstatistical data, let alone stratigraphical data, on the formsconcerned. Those data available have been plotted in Text-fig. 46C. These are based wholly on the suite of type speci-mens in each case except that Büdesheim data from Clausen(1969) are also added for early whorls. Clausen assigned thenumbered specimens as follows: C. galeatum, 36-40; C. aff.vagans, 71-75; C. acutiforme, 76-80; assignment of these nu-clei, however, is open to considerable doubt.

Considering shell form alone, Carinoceras vagans is clos-

est to C. galeatum and there is little doubt they should be re-garded as synonyms, C. vagans having priority. But the latteris here shown to modify into forms with very sharp oxyconicventers. Carinoceras acutum would seem to be close to C.acutiforme but to differ only a little from C. vagans insofaras comparisons between forms at different stages has any va-lidity. The grounds for keeping C. vagans and C. acutum sep-arate are currently based on the lack of evidence forcarination or adult growth line convexity in the former.

It is probable, as others have remarked, that Carinocerasgaleatum is a junior synonym of this group, and the larger ofthe measurements given by Wedekind (1913: 60), that areplotted on the accompanying graphs for Carinoceras, con-tinue the same trend in D/WW as the New York specimens.The stratigraphic context of the German specimens is atpresent unknown, and speculation on relationships with Eu-ropean forms is best deferred until a detailed study has beenundertaken.

Distribution.–Upper Rhinestreet Shale to middle AngolaShale (West Falls Group) in Erie andWyoming counties andprobably the equivalent interval (Grimes Siltstone to NundaSandstone (High Point Sandstone) in the Naples area (On-tario County). The stratigraphic horizon of the holotype(NYSM 3805) is unknown but the specimen almost cer-tainly came from the interval from the Grimes Siltstone tothe Nunda Sandstone of theWest Falls Group (Text-fig. 3).Clarke (1899a, b: 79) stated that the specimen came from“. . . a loose block of sandstone among the Portage outcropsin the town of Naples, . . . associated with [the brachiopods]Productella speciosa [Hall, 1867], Leptostrophia mucronata[Hall, 1867] and Amboecoelia umbonata [Conrad, 1839].”Miller (1938: 125) stated that Chadwick “. . . has recentlygiven its horizon as the Grimes sandstone member of theChemung formation” but gave no reference. Chadwick(1935a: 320-322) listed Carinoceras vagans, C. sororium, andSpheromanticoceras oxy as confined to the Chemung group(the base of which at Naples is the Grimes) but of the threeassociated brachiopods, Chadwick (1935a) listed only P. spe-ciosa as a “transient” brachiopod extending into the post-Chemung sequence. Clarke & Luther (1904) did not listany of the three brachiopods in their faunal lists of theGrimes or succeeding West Hill Flags and Sandstone (inwhich S. oxy is listed). Their faunal list for the High PointSandstone at the top of the Naples succession includes P.speciosa and A. umbonata.

We follow Pepper et al. (1956) in equating the HighPoint Sandstone at Naples with the Nunda Sandstone (Text-fig. 3) which puts the Nunda as an upper limit to the pos-sible range of the type of Carinoceras vagans.

The stratigraphically lowest specimens in our collectionsare upper Rhinestreet Shale: two specimens (NYSM 12040

Text-fig. 44. Sutures and growth line of Sphaeromanticoceras spp.from theWest Falls Group in New York and Carinoceras spp. fromNew York and Belgium. A. S. oxy (Clarke, 1897), NYSM 3747,growth line based on one of Clarke's specimens recorded as fromthe "Portage sandstone on East Hill, Naples," X 0.4. B. C. acutum(G. & F. Sandberger, 1850), IG 8633, suture at 78 mm whorlheight and reversed for comparison, of a specimen determined byMatern from the Schistes de Matagne (F3) in the railway cuttingbetween Nismes and Mariembourg, Belgium, X 0.4. C. S. oxy,NYSM 3746, the last suture, at 137 mm diameter and reversedfor comparison, of another of Clarke's specimens, from 45.7 m(150 ft) "below the Portage Sandstone on the Genesee River" (Loc.86), Wyoming and Livingston counties, X 0.4. D. C. vagans(Clarke, 1898), NYSM 12036, suture at 23 mm diameter based ona specimen from the Angola Shale (Point Breeze Goniatite Bed) onBig Sister Creek (Loc. 73/6), Angola, Erie County, X 0.8. E. S.oxy, NYSM 3748, lectotype here designated, portion of the ventralsuture on one of Clarke's specimens labeled as from Naples, X 0.4.F. S. aff. oxy, NYSM 12034, portion of a mature suture at 135mmwhorl height, reversed for comparison, showing obliquely flat-tened crest to the ventrolateral saddle, based on a specimen fromthe Relyea Creek Horizon of the Rhinestreet Shale on RelyeaCreek (Loc. 60/3), Wyoming County, X 0.4.



and 12042) from the fourth microcyclothem below the topof the Rhinestreet on Johnson Creek (Wyoming County;Loc. 58/1), and four specimens from the overlying third mi-crocyclothem down (NYSM 12048), NYSM 3200/29 fromthe Relyea Creek Horizon (Loc. 60/3), NYSM 3193 fromRelyea Creek (Wyoming County; Loc. 60/4), and NYSM3297/19 from Varysburg (Wyoming County; Loc. 57/2).

The specimens from the Angola Shale include NYSM3248/2, from the fourth microcyclothem up from the base,on Cazenovia Creek (Erie County; Loc. 75/4), and NYSM12037 from the sixth microcyclothem, just below the PointBreeze Goniatite Bed on Hampton Brook (Erie County;Loc. 74/6a). The best fauna (NYSM 12038, 12039, 12041,and 3229/6, 31) is from the Point Breeze Goniatite Bed it-self on Cazenovia Creek (Loc. 75/6). NYSM 12036 is fromthe Point Breeze Goniatite Bed on Big Sister Creek, Angola(Erie County; Loc. 73/6). The stratigraphically highest spec-imens, referred to Carinoceras aff. vegans (Pl. 23, Figs 1-4;Text-figs 43B-C) are frommiddle levels of the Angola Shale,NYSM 12043 and 12045 from 10 m above the Point BreezeGoniatite Bed at Varysburg (Wyoming County; Loc. 79-80/10), and a higher specimen (NYSM 12044) from GladeCreek (Wyoming County; Loc. 77/1).

Range.–Frasnian. Playfordites to Neomantioceras Geno-zones UD I-I-J. Regional zones of P. cf. tripartitus (22a) andSpheromanticoceras rhynchostomum (22b). Conodont ZoneMN 11-?12.

Genus DELPHICERAS Becker & House, 2000Type species.–Manticoceras unduloconstrictum Miller,

1938, by original designation (Becker &House, 1993: 114);holotype illustrated by Becker & House (1993: pl. 2, fig. 7)and herein (Pl. 18, Fig. 10).

Diagnosis:–Small-sized gephuroceratids, moderately evo-lute to subinvolute throughout ontogeny with whorl heightrather rapidly expanding. Rounded whorl section with ven-ter rounded or slightly flattened. Periodic internal shellthickenings forming constrictions on the internal molds.Growth lines rectiradiate to biconvex.

Included species.–Manticoceras unduloconstrictum Miller,1938, and Gephyroceras cataphractum Clarke, 1898.

Discussion.–The generic name originally proposed forthis group wasDelphinites Becker & House, 1993: 114, butthat proved to be preoccupied by Delphinites Sayn, 1901.Delphiceras was proposed as a replacement name (Becker &House, 2000: 140) and hence takes the same type speciesautomatically.

Distribution.–North America (Indiana and New York)and Europe (England, Belgium, Germany) (Becker &House, 1993).

Range.–Frasnian. Archoceras to Crickites Genozones UD

I-K, L.

Delphiceras cataphractum (Clarke, 1898)Pl. 19, Figs 1-9; Text-fig. 41G

Gephyroceras cataphractum Clarke, 1898, 1899a, b: 87-89, text-figs 66-67, pl. 6, figs 3-9.

Manticoceras cataphractum. Frech, 1913: 25; Miller, 1938: 77-80;House, 1962: 258; House & Kirchgasser, 1993: 275, 279.

Delphinites cataphractum. Becker & House, 1993: 114, pl. 2, figs4-6.

Delphiceras cataphractum. Becker & House, 2000: 130, 140.

Type material.–Seven original cotypes of Clarke (NYSM3638-3644; formerly 12160/1-7). Lectotype here chosen asNYSM 3643 (figured by Miller, 1938: pl. 13, fig. 1 andherein Pl. 19, Fig. 1 and Text-fig. 41G). These fall into twogroups: NYSM 3638 and 3643 are limestone internalmolds; NYSM 3639-3642 and 3644 are baritic shell replace-ments.

Description.–Detailed descriptions have been given byClarke (1899a, b: 87-89) andMiller (1938: 77-80) and neednot be repeated here. The prominent constrictions are diag-nostic.

Distribution.–Lower Hanover Shale (West Falls Group;Wyoming and Chautauqua counties). The cotypes arerecorded as from calcareous concretions near the village ofJava (Wyoming County), at levels subsequently referred tothe Hanover Shale; the horizon at Java is in Beaver MeadowCreek (Bed 92/2). We have only found specimens in thenodules in the lowermost microcycles in the Hanover Shale,including localities close to Lake Erie Shore (Text-fig. 16),such as Walnut Creek (Loc. 89a/2-4), south of Silver Creek(Chautauqua County).

Range.–Frasnian: Archoceras Genozone UD I-?K. Re-gional Zone of Delphiceras cataphractum (23). MN 12.

Genus PLAYFORDITES Becker, House &Kirchgasser, 1993

Type species.–Goniatites lamed var. tripartitus G. & F.Sandberger, 1850. Original material lost; neotype chosen byBecker & House (1993: 121) is the specimen figured byHouse (in House & Ziegler, 1977: pl. 3, figs 13-15), andby Becker et al. (1993: fig. 4, O, P; Univ. Marburg Coll.,Mbg. 2594a).

Diagnosis.–Slender gephuroceratids with discoidal shellin middle whorls with spiral thickenings on either side ofventer or on flanks, which are not shown on external surfaceof shell.

Included species.–Playfordites tripartitus (G. & F. Sand-berger, 1850).

Distribution.–North America (New York), Europe (Ger-

many), and Western Australia.Range.–Frasnian: Playfordites to Neomanticoceras Geno-

zones UD I-I-J.

Playfordites cf. tripartitus (G. & F. Sandberger, 1850)Text-fig. 42A; Table 17

Playfordites cf. tripartitus. House & Kirchgasser 1993: 276.

Material.–Two small specimens only (NYSM 12074-12075), both preserved in mudrock and calcite replacementas internal molds showing part of the body chamber andphragmocone.

Dimensions.–See Table 17.Description.–Subinvolute, with open umbilicus and well-

rounded shoulders (Text-fig. 42A) with maximum widthclose to shoulder and flanks converging convexly towardrounded venter. In NYSM 12074, ventrolateral furrows arewell developed with arched band between. In NYSM12075, in addition to ventrolateral groove, is another butslighter spiral concavity on outer flanks, umbilicad of fur-row. Growth lines forming slight salient on umbilical shoul-der and sinus, which centers low on flanks; lines thensweeping well forward to ventrolateral salient on umbilicalside of furrow; ventral sinus deep, rounded. Suture (NYSM12074) showing small, undivided ventral saddle, broadlyrounded and nearly symmetrical lateral saddle, and broadand rounded lateral lobe (L) low on flanks. Ventral wrinklelayer showing as fine, discontinuous striae on internal mold,which are directed in convex rursiradiate course acrossflanks.

Discussion.–This is a very rare form and so, unfortu-nately, comparison cannot be made with the varied develop-ments of specimens from the Canning Basin, Australia.

Distribution.–Upper Rhinestreet Shale to Lower AngolaShale (West Falls Group) in Wyoming County. NYSM12074 from the Upper Rhinestreet Shale, Johnson Creek(Loc. 58/1) and NYSM 12075 from the Point Breeze Goni-atite Bed of the Angola Shale, Relyea Creek (Loc. 82/6).

Genus SPHAEROMANTICOCERAS Clausen, 1971Type species.–Goniatites affinis Steininger, 1849, by orig-

inal designation (Clausen, 1971: 197).

Diagnosis.–Small to very large gephuroceratids withwidely umbilicate earliest whorls followed by stages with avery depressed whorl section, often banana-shaped. Outerwhorls of large forms laterally compressed and discoidal. Su-ture with mid-lateral saddle very high, symmetric to asym-metric with steep to overhanging dorsad face. Growth linesbiconvex in early stages and small forms to subbiconvex tosubconvex in very large discoidal specimens. Wrinkle layercharacteristically with much finer striae (in type species)than in Manticoceras.

Included species.–Goniatites affinis Steininger (1849, nom.nud.; 1853: 42, pl. 1, figs 4, 4c);Manticoceras hunanense Xu,1977; M. kwangsiense (Chao, 1956: 106, pl. 1, figs 19-22;= affine fide Becker &House, 1993);M. kweipingenseChao,1956 (? = orbiculus fide Becker in litt.); Goniatites lamed var.latidorsalisG. & F. Sandberger (1850: 90, pl. 8, fig. 8; = affi-nis fide Clausen, 1969); Ammonites orbiculus Beyrich (1837:36, pl. 2, figs 4a-b); M. oxy Clarke, 1897; A. primordialisvon Buch (1832: 168, pl. 1, figs 15-17; possible senior syn-onym of affinis);M. rhynchostomum Clarke, 1898; Sphaero-manticoceras rickardi House & Kirchgasser, 1993.

Discussion.–Although it is accepted that Clausen’s taxo-nomic separation of this group is useful, so many of the typespecimens of species that he included within it are poorlyknown and badly described that there is more than usualuncertainty on the assignments given here. A further prob-lem is that Sphaeromanticoceras affine (Steininger, 1849: 2-3, 42, figs 4, 4a), the type species, is very small(approximately 11 mm in diameter), whereas late Frasnianspecies can reach over a third of a meter in diameter.

Distribution.–Widely distributed in North America, Eu-rope, Asia, and Australia.

Range.–Frasnian: Prochorites to CrickitesGenozones UDI-F-L.

Sphaeromanticoceras rhynchostomum (Clarke, 1898)Pl. 20, Figs 1-12, Pl. 21, Figs 1-12, Pl. 22, Figs 1-9; Text-figs

41F, 42B, H, I, 43F-G, 47; Table 18

Manticoceras rhynchostoma Clarke, 1898, 1899a, b: 65-69, 81, pl.4, figs 6-13, pl. 5, fig. 1, text-figs 30-43; 1904: 356, 361, 365;Frech, 1913: 24; Miller, 1938: 99-102, pl. 14, fig. 19, pl. 25,figs 1-8, pl. 26, figs 1-2, text-figs 18A-L; Clausen, 1969: 146.

Table 17. Biometric data for Playfordites cf. tripartitus (G. & F. Sandberger, 1850) from the Rhinestreet Shale.


NYSM 12074 9.2 4.4 4.4 ca. 2.0 1.00 0.225.6 3.1 2.6 1.5 1.19 0.273.7 2.1 1.5 1.2 1.40 0.32

NYSM 12075 8.2 4.3 — — — —


Sphaeromanticoceras rhynchostomum. House & Kirchgasser, 1993:275-276, figs 4I-J.

Type material.–Cotypes are NYSM 3784-3796, and ofthese, NYSM 3796 is here designated as lectotype (figuredby Clarke, 1898, 1899a, b: 68, text-fig. 43, and herein, Pl.22, Figs 6-7, Text-fig. 47A). All are reported from Big SisterCreek (Erie County), and probably from the Angola Shaleat Angola (Erie County; Loc. 73).

New Material.–One-hundred fifteen specimens (includ-ing NYSM 12035, 12061, and 12080-12093. Of these, 40are from Rhinestreet Shale, 15 from the Angola Shale belowthe Point Breeze Goniatite Bed, 43 from the Point BreezeGoniatite Bed or adjacent beds (NYSM 12087-12093), and7 from the Angola Shale equivalents above the Point BreezeGoniatite Bed.

From the Rhinestreet Shale in the Relyea Creek Horizonor nearly correlative horizons: NYSM 12084/1-4, CazenoviaCreek (Erie County; Loc. 54/1); NYSM 12082, Varysburg(Loc. 57/1); NYSM 12085, Johnson Creek (Loc. 58/1); sev-eral specimens in collection 3200, Relyea Creek (Loc. 60/3);NYSM 12080/1-6, Stony Creek (Loc. 61/1); and NYSM12081/1, 2, Kennedy Gulf (Wyoming County; Loc. 63/3).Included is a fine collection (NYSM 6680/A-H, J-Q, and12132) by L. V. Rickard from the vicinity of the RelyeaCreek Horizon at Relyea Creek (Loc. 60).

From the Angola Shale, the best material is from thePoint Breeze Goniatite Bed: NYSM 12087/1-4, PointBreeze (Lake Erie Shore) (Loc. 72/6); NYSM 12088/1-3,Big Sister Creek, Angola (Loc. 73/6); NYSM 12035, 12089,and 12090/1-6, bed below the Point Breeze Goniatite Bed

(Loc. 74/6a); NYSM 12078 and 12079, Hampton Brook(Loc. 74/6b) in Erie County; and NYSM 12091/1-4,Cazenovia Creek (Loc. 75/6); NYSM 12092, Johnson Creek(Loc. 81/6); NYSM 12093/1-23, Relyea Creek (Loc. 82/6);and coll. #3205, Stony Creek (Loc. 83/6a), in WyomingCounty.

From higher in the equivalents of the Angola Shale,Sphaeromanticoceras rhynchostomum (or S. oxy) can be repre-sented in the vicinity of the Nunda Sandstone at Varysburg(Gassman Road) (Loc. 80/14) and at Wolf Creek (Loc. 85)below Castile in Wyoming County.

Dimensions.–Dimensions are given in Table 18. Accom-panying graphs separate type specimens (Text-fig. 47A), andspecimens from the Rhinestreet (Text-fig. 47B) and Angolashales (Text-fig. 47C).

Description.–Full descriptions were given by Clarke(1898, 1899a, b) and repeated by Miller (1938). Some ma-terial is in the form of baritic replacements and some typematerial consists of gutta percha casts. The bulk of the newmaterial is preserved as calcitic replacements with calcitic ormudrock infills of phragmocone chambers, whereas thebody chambers are mostly preserved in calcareous mudrock.

Protoconch approximately 0.8 mm in diameter andmaximum width. (That the protoconch is wider than thefirst whorl is clear from NYSM 12079; Pl. 20, Fig. 4.) Nolirae on the protoconch (ammonitella), which has a diameterof approximately 1.1 mm. (An eccentrically placed am-monitella is shown in Pl. 22, Fig. 9.)

Whorl following ammonitella widely umbilicate, withperiodic raised lirae, numbering approximately 17 per whorland almost rectiradiate across flanks and venter (Pl. 20, Figs

Table 18. Biometric data for Sphaeromanticoceras rhynchostomum (Clarke, 1898) from the upper Rhinestreet and lower Angola shales. SeeText-fig. 47 for graphs.


NYSM 3796 94 32 52 ca. 13 0.62 0.13(lectotype)NYSM 12132 27.9 16.0 15.7 7.0 1.02 0.25NYSM 12080 31.1 15.3 15.0 8.6 1.02 0.28NYSM 12085 17.2 10.5 7.4 5.4 1.42 0.31

11.3 7.1 4.5 4.0 1.58 0.357.9 5.0 2.9 3.1 1.72 0.395.4 3.3 1.8 2.2 1.83 0.414.0 2.3 1.2 1.8 1.92 0.452.9 1.6 0.9 1.3 1.78 0.452.2 1.2 0.7 0.9 1.71 0.411.6 0.8 0.5 0.7 1.60 0.44

NYSM 12088a 13.2 8.3 5.7 4.2 1.46 0.458.6 5.9 3.2 3.4 1.84 0.406.1 4.2 1.9 2.8 2.21 0.344.6 2.8 1.3 2.0 2.15 0.433.3 1.9 1.1 1.5 1.73 0.452.5 1.3 0.7 1.1 1.86 0.44


2-3).Whorl width increasing progressively to produce tubbyshells by 10 mm diameter (Pl. 21, Figs 1-12, Pl. 22, Fig. 9;Text-figs 42B, H-I). Thereafter, WW/WH ratio declining(Text-figs 47A-C), reversing earlier pattern; change canoccur earlier in Rhinestreet than in Angola material. Duringthis stage, pattern of earlier lirae continuing but with gradualonset of slight biconvexity, increasing in frequency to ap-proximately 40 per whorl, with stronger development onumbilical wall and lower flanks. Thereafter ornament pro-gressively lost. Early lirae very dissimilar to virtual flares inearly stages of Manticoceras sinuosum Group.

Subsequent growth as described by Clarke and Miller.Lectotype (Pl. 22, Figs 6-7; Text-fig. 47A) shows well thegradual change inWW/WH ratio and how shells above ap-proximately 35 mm diameter become laterally compressed.

Because of preservation, sutural ontogeny of earlieststages not determined. Lectotype with relatively high me-dian saddle (MSh/ELh = 0.75 at 85 mm diameter) at whichpoint, for slightly asymmetric lateral saddle, LSw/LSh = 1.0.Lobe low on flanks (L) already as sharp as at 20 mm in somespecimens. Note that in NYSM 12035, at WH = 61 mm(Text-figs 43F-G), the median saddle is very high (MSh/LSh= 1.14) and the low lateral lobe (L) very sharp.

Discussion.–It is the characteristic tubby form in stages toapproximately 20 mm diameter and the distinctive early li-rate ornament that distinguishes this species. It is clear thatlarge specimens become increasingly laterally compressedand discus shaped. End forms of this line are here placed inSphaeromanticoceras oxy. A case could be made for including

these within S. rhynchostomum as is discussed below.Distribution.–Upper Rhinestreet Shale and Angola Shale

(West Falls Group) in Erie and Wyoming counties. Com-mon in Relyea Creek Horizon of the upper RhinestreetShale in Wyoming and Erie counties and the Point BreezeGoniatite Bed of the lower Angola Shale in Erie County.

Range.–Frasnian: Playfordites andNeomanticocerasGeno-zones UD I-I to J. Regional Zone of P. cf. tripartitus (22a)to Sphaeromanticoceras rhynchostomum (22b). ConodontZone MN 11-?12.

Sphaeromanticoceras oxy (Clarke, 1897)Text-figs 44A, C, E, 45, 46B; Table 19

Manticoceras oxy Clarke, 1897: 53; 1898, 1899a, b: 53, 77-79,82, text-fig. 61, pl. 2, figs 5-6, pl. 3, figs 1-3 (not fig. 4); Frech,1913: 24; Miller, 1938: 93-95, text-fig. 17, pl. 16, fig. 1, pl.17, fig. 1, pl. 18, figs 1-3.

Carinoceras oxy. House & Kirchgasser, 1993: 276.

Type material.–The cotypes have been described and il-lustrated by Clarke and by Miller and comprise the follow-ing figured by Clarke, 1898, 1899a,b: pl. 2, figs 5-6,(NYSM 3746, a fine but crushed specimen), pl. 3, fig. 1(NYSM 3747, growth line illustrated in Text-fig. 44A), pl.3, figs 2-3 (NYSM 3748, lectotype here designated, suturerefigured inText-fig. 44E) and text-fig. 61 (NYSM 12305/4,another large crushed specimen in which the whorl outlinecannot be determined). None of these large and impressivespecimens shows inner whorls, and the largest specimens

Table 19. Biometric data for Sphaeromanticoceras oxy (Clarke, 1897) from the Rhinestreet and Angola shales. See Text-fig. 46 B for graph.


NYSM 3746 201 — 105 24 — 0.12(cotype)

NYSM 3747 — ca. 47 ca. 150 — — —(cotype)

NYSM 12034 355 — 177 ca. 32 — 0.09NYSM 16536 299 ca. 100 162 39 0.61 0.13

176 ca. 70 75 29 0.93 0.16147.7 52.4 81.2 19.3 0.65 0.13101 47 50 23 0.94 0.2387.4 31.8 46.0 15.2 0.69 0.1761 33 28 16 1.18 0.2648.8 23.1 24.1 11.8 0.96 0.2429.7 19.2 13.6 7.4 1.41 0.2519.3 11.4 7.7 6.2 1.48 0.3212.9 7.8 5.3 4.6 1.47 0.368.7 5.2 3.0 3.5 1.73 0.406.8 4.3 2.2 2.5 1.95 0.375.0 3.4 1.5 1.9 2.27 0.38

NYSM 12046 ca. 137 — ca. 74 ca. 13 — 0.06NYSM E-355 125 20 74 ca. 8.0 0.27 0.10NYSM 12047 16.6 6.2 8.1 2.8 0.77 0.17


Text-fig. 45. Sutures and cross sections of Sphaeromanticoceras oxy (Clarke. 1897). A. NYSM 16536, cross section and suture based on a spec-imen collected by J. Kralick from the lower Angola Shale of Kennedy Gulf (Loc. 63/4), south of Dale, Wyoming County. B. NYSM 16569,cross section and suture based on a specimen collected by L. V. Rickard near the Relyea Creek Horizon, Upper Rhinestreet Shale, RelyeaCreek (Loc 60/3), Wyoming County. C-D. NYSM 16570, side view and partial cross section of a specimen from a loose block, collectedby B. Oldfield, of a shell-rich channel fill, upper West Falls Group (Nunda or Wiscoy Sandstone equivalents), at Loc. 87a, US Rte. I-390in Steuben County, southeast of Dansville, Livingston County. Scales = 5 cm.


Text-fig.46.Graphsshowingtheontogenyof

Carinocerasand

Sphaerom

anticoceras.A.C

.vagan

s(Clarke,1898)Group,basedonmaterialfromtheRhinestreetandAngolashales;

seeTable16forbiometricdata.B.S

.oxy(Clarke,1897)[asC.oxy],basedonmaterialfromtheRhinestreetandAngolashales;seeTable19forbiometricdata.C.C

.spp.,based

onmaterialfromGermanyandRussiaasindicated.


Text-fig.47.Graphsshowingtheontogenyof

Sphaerom

anticoceras

rhynchostomum(Clarke,1898)from

theRhinestreetandAngolashalesofwesternNewYorkState;seeTable18

forselectedbiometricdata.A.Basedoncrosssections(correctedforsize)ofNYSM

3796,thelectotypeheredesignated,figuredbyClarke(1899a,b:text-fig.43)andNYSM

3794

(figuredbyClarke,1899a,b:text-fig.41).Bothprobablyfrom

thelowerAngolaShalefrom

eitherBigSisterCreekorAngola,ErieCounty.B.Datafrom

23specimensfrom

the

upperRhinestreetShale(Locs.54/1,57/1,57/2,58/1,60/3,61/1,and63/3),allwithin3-10m(10-33ft)belowthetopatlocalitiesbetweenCazenoviaCreek,ErieCounty,and

KennedyGulf,WyomingCounty;NYSM

12080-12086.C.Databasedon30specimensfrom

thelevelofthePointBreezeGoniatiteBedofthelowerAngolaShale(Locs.72/6,

73/6,74/6a,75/6,81/6,82/6,and83/6a)betweenPointBreeze,LakeErieshoreandStonyCreek,WyomingCounty;materialinNYSM.



have their discoidal form enhanced by lateral crushing, alto-gether an unsatisfactory combination. In selecting NYSM3748 as lectotype, emphasis has been on the solid preserva-tion indicating a rounded, rather than sharp, venter, andhence enabling comparison with the better quality materialdescribed below.

Additional material.–The finest new material is a speci-men collected by J. Kralick (NYSM 16536) from the lowerAngola Shale at Kennedy Gulf (Loc. 63/4). Ten specimenscollected in this investigation are included but only NYSM12034 (as Sphaeromanticoceras aff. oxy, Text-fig. 44F) and12046-12047, 12133, 16569, and 16570 with confidence.

Dimensions.–See Table 19.Description.–The interpretation of this large species has

depended on the availability of material showing not onlythe large discoidal outer whorls but also the earliest stages(NYSM 16536) that are essentially those of Sphaeromantic-oceras rhynchostomum. This accords with the very large spec-imen specimen figured by Clarke (andMiller, 1938: text-fig.17) and the lectotype here designated.The umbilical width continues to be small with growth.

Most specimens show some degree of crushing, but the max-imum whorl width continues to be near the shoulder andthe flanks converge flatly to the venter. Clarke’s restored out-line (NYSM 12305/4) shows a carinate venter, but the in-ternal molds of other specimens are often narrowly roundedrather than acute, but this could reflect loss of the test; suchis the case in the largest specimen seen (NYSM 12034). Su-tures of several of the larger specimens are illustrated here(Text-figs 44 C, E-F) and it will be noted that the ventrolat-eral saddle can have a flater ventrad face (NYSM 12034:Sphaeromanticoceras aff. oxy) or can be more symmetrical.Several specimens show adult growth lines and these areconvex (NYSM 3747) with a very prominent ventrolateralsalient and what was formerly a latero-umbilical salient nowbeing only an undulation in the growth line course low onthe flanks (Text-fig. 44A).

Discussion.–The source of the Sphaeromanticoceras oxygroup is now clearly indicated as from S. rhynchostomum.Because the range is similar, it raises the question of whetherthe former is the macroconch form and the latter a micro-conch and they represent a dimorphic pair. Further com-ment on this would require elucidation of the number ofwhorls in each, and there is insufficient material available toattempt this.

Distribution.–Middle Rhinestreet and Angola shales(Point Breeze Goniatite Bed) and equivalents (GardeauShale and West Hill Shale and Sandstone) (West FallsGroup) in Erie, Wyoming, Livingston, Steuben, and On-tario counties, and Nunda orWiscoy sandstones (West FallsGroup) in Wyoming, Livingston, and Steuben counties.

Of Clarke’s cotypes, NYSM 3746 came from 45.7 m(150 ft) “below the Portage sandstone, at the Lower PortageFalls on the Genesee River” and Clarke recorded that thiswas the earliest record known to him. This locality (GeneseeGorge, Loc. 86, Wyoming and Livingston counties) is fig-ured in Text-fig. 12 but the exact level of Clarke’s specimenis unknown; on our correlation (Text-fig. 3) the Table RockSandstone in the Gardeau Shale at the top of the Lower Fallsis equivalent to a level high in the Rhinestreet Shale of ErieCounty, not to the Angola Shale as Clarke supposed. NYSM3747 from East Hill and NYSM 3748 (the lectotype desig-natd here) from Caulkins Gully, Naples (Ontario County),probably came from the Gardeau Shale and also NYSME355, reported from the West Hill flags in the village ofNaples referred to by Miller (1938: 95).The new material from the Rhinestreet Shale is wholly

from the upper part of the unit. The collection from the Re-lyea Creek Horizon (Loc. 60/3), and levels in the meterbelow, at Relyea Creek (Wyoming County; Loc. 60), includethe large specimen NYSM 12034 (Sphaeromanticoceras aff.oxy) and NYSM 16569 collected by L. V. Rickard.The best new material from the Angola Shale is NYSM

16536 from the lower Angola Shale at Kennedy Gulf(Wyoming County; Loc. 63/4; Text-fig. 13) and from thePoint Breeze Goniatite Bed, NYSM 12046 and 3245/2, BigSister Creek (Loc. 73/5), NYSM 3229/2, 11, CazenoviaCreek (Loc. 75/6), and NYSM 12047, the bed below thePoint Breeze Goniatite Bed, Hampton Brook (Loc. 74/6a)in Erie County. NYSM 12133 is from the eighth microcy-clothem of the Angola Shale in Stony Creek (Loc. 83/8) inWyoming County.Higher records in the upper West Falls Group include

Clarke’s 320-mm-diameter specimen from Stony BrookGlen, Dansville (Steuben County; Loc. 87), mentioned byMiller (1938: 95); this specimen could be from Angola Shaleequivalents because we consider the Scraggy Bed level mark-ing the base of the Angola to lie low in the section there.The Dansville area is also the source of NYSM 16570 fromthe Nunda Sandstone or Wiscoy Sandstone equivalents inthe roadcut I-390-Dansville (Steuben County; Loc. 87a).Also reported from the Wiscoy Sandstone is Clarke’s giantspecimen (460-600 mm diameter) from Castile (WyomingCounty; Miller, 1938: 95). The possibility that the WiscoySandstone records could be the eastern representative ofSpheromanticoceras rickardi cannot be discounted.

Range.–Frasnian: Playfordites andNeomanticocerasGeno-zones UD I-I-J. Regional Zone of P. cf. tripartitus (22a) toSphaeromanticoceras rhynchostomum (22b). Conodont ZoneMN 11-?12.

Sphaeromanticoceras rickardi House & Kirchgasser, 1993Text-figs 48A-C; Table 20

Crickites holzapfeli (Wedekind, 1913). Matern, 1931b: text-figs1a-b; House, 1978: 62.

Sphaeromanticoceras rickardiHouse & Kirchgasser, 1993: 274, figs5D-F.

Type material.–The species is based on IG 4449, a spec-imen from 560 m southeast of Romerée, chemin deMatagne-la-Petite, Belgium, and figured by Matern (1931b:text-figs 1a-b).

New York material.–Only two specimens are known, alarge, partly crushed, phragmocone (NYSM 12032) and asolid, incomplete specimen (NYSM 12033) showing septaand approximately half a whorl of body chamber, both fromthe Hanover Shale.

Dimensions.–See Table 20.Description.–Phragmocone of NYSM 12032 involute,

with small, deep umbilicus, robust whorl section with max-imum width low on flanks that curve to rounded shoulder.Only small part of body chamber preserved. Growth linesnot seen. Suture with distinctive asymmetrical lateral saddlewith nearly vertical dorsad face (LSh/LSw =1.17; Text-fig.48C). Smaller specimen (NYSM 12033) showing half whorlof body chamber and adjacent proximal phragmoconeshowing involute form, deep umbilicus, and maximumwhorl width low on flanks that converge convexly to well-rounded venter. Complete suture not seen; growth lines in-distinct.The specimen in the collection of G. Kloc (University of

Rochester) with a lateral saddle with a less steep dorsad faceto the lateral saddle was referred by us to Sphaeromanticocerasaff. rickardi (see House & Kirchgasser, 1993: figs 5G-1.

Discussion.–Growth lines are not shown on these speci-mens so their earlier assignment to Crickites (Kirchgasser &House, 1981: 45, text-fig. 3) cannot be justified. Further,subsequent work on that genus has shown that the rotundwhorl sections of the C. holzapfeli and C. lindneri Groupcontinues through ontogeny to a large size. By contrast,Sphaeromanticoceras rickardi becomes more compressed inlater stages. In his revision of late Frasnian forms from Bel-gium, Matern (1931b) illustrated type material. He arguedthat Wedekind’s type material of C. holzapfeli, being lost, a

neotype should be erected. He proposed for this a specimenfrom Belgium, rather than from the German type area.

Table 20. Biometric data for Sphaeromanticoceras rickardi House & Kirchgasser, 1993, from the Hanover Shale.


NYSM 12032 209 max.140 60 (est.) 88 ca. 15 0.68 0.11

NYSM 12033 130 max.92 47 52 — ca. 0.90 —

Text-fig. 48. Sutures and cross sections of Sphaeromanticoceras,Crickites, and ?Archoceras. A-C. S. rickardi House & Kirchgasser,1993, from the Hanover Shale in New York and from Belgium. A.IG 4916, suture at 165 mm diameter based on a specimen from2,700 m east-northeast of Roly (Sautour Sheet), Chemin deViller's en Fagne, X 0.6. B. IG 4449, the "Neotype" figured byMatern (1931b: text-fig. 1a-b) from 560 m south-southeast ofRomerée, Chemin de Matagne-la-Petite (Surice Sheet) at about100 mm diameter (suture for comparison), X 0.6. C. NYSM12032, suture of a specimen at about 137 mm diameter from theHanover Shale collected by L. V. Rickard at Glade Creek (Loc.91/7), Strykersville, Wyoming County, X 0.6. D-F. C. holzapfeliWedekind, 1913, whorl sections and a suture based onWedekind'scotypes from Bicken, Germany (afterWedekind, 1913). Originalsassumed to be natural size, X 0.6. G. ?Archoceras sp., NYSM16587, cross section of a specimen from a concretion in theHanover Shale, Erie County, 4.7 m below the base of the DunkirkShale in Irish Gulf (Loc. 90). H. ?Crickites sp. juv., NYSM 16589,from the same locality and horizon in Irish Gulf (Loc. 90). Scales= 1 mm (G-H).


Some ofWedekind’s type material however, appears to havesurvived, so this assignment is invalid. The suture ofMatern’s “neotype” (IG4449) from the Schistes de Matagne(Text-fig. 48B), although not the same size as the New Yorkspecimen, shows a similar shape, particularly in the upright,asymmetrical lateral saddle at D = 100, LSh/LSw = 1.26. Ina rather larger specimen (IG4916) from near Roly (Text-fig.48A), at D = 165, LSh/LSw = 1.24. The height of the me-dian saddle in this species is somewhat variable.

Distribution.–Type specimens, Europe, Matagne Shaleof Belgium, and North America (New York), Hanover Shale(Wyoming County).NYSM 12032, found by L. V. Rickard, from upper

Hanover Shale, Glade Creek, Strykersville (WyomingCounty; Loc. 91/7). NYSM 12033 was found loose justabove the base of the Hanover Shale also in Glade Creek(Loc. 91). The G. Kloc specimen of Sphaeromanticoceras aff.rickardi came from 1.5 m above the base of the HanoverShale in Walnut Creek (Cattatraugus County; Loc. 89a/5).

Range.–Frasnian: NYSM 12032: Crickites GenozoneUD I-L, Regional Zone of Sphaeromanticoceras rickardi(24b). Conodont Zone MN 12-?13. The type specimensfrom Belgium from Archoceras Genozone UD I-K. LowerHanover specimens: NYSM 12033 and Kloc specimen ofS. aff. rickardi: Crickites Genozone UD I-L, Regional Zoneof C. lindneri (24a). Conodont Zone MN 12.

Subfamily CRICKITINAEWedekind, 1913Genus CRICKITESWedekind, 1913

Type species.–Crickites holzapfeli Wedekind (1913: 72),by subsequent designation (Wedekind, 1917: 130).

Diagnosis.–Subinvolute, medium- to large-sized manic-oceratids with convex growth lines from earliest stages withsinus on umbilical seam and broad salient over flanks.Whorl section distinctively rounded (rotund) to rounded-rectilinear with whorls converging convexly to rounded ven-ter (WW/WH = 1.0-1.3). Suture as inManticoceras.

Included species.–Genus restricted here to forms withspecified whorl outline: Crickites holzapfeliWedekind (1913:72, pl. 7, figs 5-6, text-figs 14a-c); ?Manticoceras cordiformeMiller [1938: 82, pl. 21, figs 2-4; = ?M. septentrionaleMiller,1938: 102, pl. 23, figs 1-2, subjective), a more laterally com-pressed form and in the adult (fide House & Pedder, 1963,pl. 76, figs 10-11) with a very slight sinus on the lateralsalient]; M. lindneri Glenister [1958: 72, pl. 6, figs 1-2(paratype), pl. 7, fig. 2, pl. 9, figs 1-2 (holotype), text-fig. 4,5B, C; one of the syntypes (UWA 35774) has a slight sinuson the lateral salient]; and M. neverovi Bogoslovsky (1958,pl. 3, fig. 1). Forms with very high lateral saddles, such asM.altaicum Bogoslovsky (1958: 100, pl. 4, fig. 1, text-fig. 25)should, perhaps, be separately placed.

Discussion.–Matern (1931b) created a Belgian neotypefor Crickites holzapfeli, but this is invalid because the originaltypes were then still available in Göttingen but they mightsubsequently have been destroyed during World War II. Itseems best to base the species on the original figures of thetype specimens from Adorf, Germany, figured byWedekind(1913; refigured here, Text-figs 48D-F).The later whorls of some gephuroceratids show convex

growth lines and ifWedekind’s subfamily is retained, it mustrest on the presence of convex growth lines from the earlieststages; these are known from approximately 9 mm diameterin the type species (House, 1963: pl. 3, fig. a).

Distribution.–Widely distributed. North America (west-ern Canada, New York), Europe, Russia, North Africa, andWestern Australia.

Range.–Frasnian: CrickitesGenozone UD I-L. ConodontZone MN 12-13.

Crickites lindneri (Glenister, 1958)Text-fig. 49; Table 21

Manticoceras lindneriGlenister, 1958: 72, pl. 6, figs 1-2, pl. 7, fig.2, pl. 9, figs 1-2, text-figs 4, 5B-C.

Sphaeromanticoceras lindneri. Becker, House & Kirchgasser, 1993:

Table 21. Biometric data for Crickites lindneri (Glenister, 1958) from the Virgin Hills Formation (UWA), Western Australia, and theHanover Shale, New York.

Specimen D WW WH-Wh UW WW/WH UW/D

UWA 34765 195 85 — — — —(holotype)

UWA 35766 209 89 98-76 ca. 37 0.91 0.18(paratype) 131 74 67-ca. 55 25 1.10 0.19

74 50 40-31 ca.15 1.25 0.20MBC 1983 — 94 ca. 75- — — 1.25 —NYSM 16537 107 59.8 54-41.3 ca. 19.9 1.11 0.19

66 ca. 42.5 ca. 33.5-23.6 ca. 12.4 1.26 0.19ca. 39.6 28.1 20.9-15.2 — 1.34 —

NYSM 16538 — 62.8 47.9- — — — —


Text-fig. 49. Sutures and cross sections of Crickites lindneri (Glenister, 1958) based on specimens from the lower Hanover Shale on WalnutCreek (Loc. 89a), Silver Creek, Chautauqua County. A-B. NYSM 16537, cross section and suture of a specimen collected by G. Klapperabout 100 yd (93 m) upstream of the NY Rte. 20 bridge, X 0.9. C. NYSM 16538, suture based on a loose specimen collected by G. Klapperfrom a similar area to the preceding, X 0.8. D-E. HM MBC 1943, growth line and cross section based on a specimen collected by R. T.Becker, from the same area, X 0.8. Scales = 5 cm.


301-314.

Type material.–The type specimens of Glenister (holo-type, UWA 34765) are in the collections of the Universityof Western Australia, Perth, where we have examined them.

New material.–Two loose specimens (NYSM 16537 and16538) collected by G. Klapper from the lower HanoverShale onWalnut Creek (Loc. 89a); one loose specimen (HMMBC 1943) collected by R. T. Becker from the same local-ity; a fine specimen shown to us by G. Kloc from approxi-mately Bed 89a/5 of the same section (Text-fig. 16).

Dimensions.–See Table 21.Description.–NYSM 16537 showing cross section indi-

cating development of subinvolute specimen with rotundwhorl section from at least 2 cm diameter to maximum seenof over 10 cm (Text-fig. 49A). Suture typically gephurocer-atid but with high, divided median saddle (Text-fig. 49B)but rather symmetrical lateral saddle. NYSM 16538 (frag-ment of larger specimen) showing suture with less high me-dian divided saddle and more asymmetrical lateral saddle(Text-fig. 49C). Fragment of even larger specimen (HMMBC 1943) showing subcircular cross section and growth

Table 22. Biometric data for Probeloceras lutheri (Clarke, 1885) from the Cashaqua shale. See Text-fig. 35C for graph.


NYSM 4067 ca. 29.6 — ca. 10.0 ca. 12.2 — 0.41NYSM 13736 40.3 — 11.2 20.2 — 0.50NYSM 12721 33.0 — 9.8 15.2 — 0.46NYSM 12739 29.5 — 9.6 13.0 — 0.44NYSM 12720 29.1 — 9.0 13.1 — 0.45NYSM 12175 ca. 27.8 — 8.5 12.1 — 0.44

20.8 — 7.1 8.4 — 0.4014.9 2.9 5.3 5.9 0.55 0.4010.6 — 3.8 4.3 — 0.417.5 1.8 2.6 3.1 0.69 0.414.0 1.1 1.3 1.8 0.85 0.452.9 0.90 0.90 1.3 1.00 0.452.1 0.70 0.65 0.95 1.10 0.45

NYSM 12722 26.0 — 8.5 11.1 — 0.43NYSM 12741 23.2 4.5 7.6 9.7 0.59 0.42

12.5 2.6 4.2 5.3 0.62 0.429.2 2.0 3.0 3.9 0.67 0.426.7 1.7 2.3 2.8 0.74 0.424.8 1.3 1.6 1.9 0.81 0.403.5 1.1 1.2 1.3 0.92 0.37

NYSM 12742 21.5 4.3 8.0 8.8 0.54 0.41NYSM 12740 17.2 — 5.6 7.1 — 0.41NYSM 3648 17.2 — 6.5 6.4 — 0.37NYSM 12737 15.5 ca. 3.0 4.9 6.9 0.61 0.45

11.6 2.5 3.8 4.9 0.66 0.428.4 2.1 2.9 3.5 0.72 0.426.0 1.6 2.0 2.5 0.80 0.424.3 1.2 1.5 1.8 0.80 0.423.0 0.95 0.97 1.3 0.98 0.432.1 0.75 0.69 0.93 1.09 0.44

NYSM 12723 13.8 3.3 5.5 4.7 0.60 0.349.4 2.4 3.6 3.5 0.67 0.37

NYSM 12744 7.3 1.9 2.4 3.1 0.79 0.425.3 1.5 1.8 2.3 0.83 0.43

Text-fig. 50. Sutures and cross sections of Probeloceras lutheri (Clarke, 1885) from the Cashaqua Shale, New York, mainly based on Kirchgasser(1975). A-G. From the lower Cashaqua Shale, Cayuga Creek (Loc. 33/5), Cowlesville, Wyoming County. A. NYSM 12727, suture and crosssection at 1.5 mm diameter, X ca. 24. B. NYSM 12729, reversed suture at ca. 2.7 mm diameter, X 24. C. NYSM 12728, suture and crosssection at 3.9 mm diameter (2.75 whorls), X 12. D. NYSM 12729 reversed suture at 7.6 mm diameter, X 12. E. NYSM 12734, reversedsuture at ca. 10.0 mm diameter (ca 4.25 whorls), X 12. F. NYSM 12738, sutures at ca. 26 mm diameter and six septa previous and crosssection, X 4. G. NYSM 12736, suture at 39.4 mm diameter (ca 6.5 whorls), X 4. H. NYSM 12744, cross section at 7.3 mm diameter basedon a specimen from the Parrish Limestone, Conklin Gully (Loc. 47/2), near Naples, Ontario County, X 4. I. NYSM 12737 (continued right)


(Text-fig. 50 continued) cross section at 15.4 mm diameter, from the lower Cashaqua Shale, Cayuga Creek (Loc. 33/5), Cowlesville,Wyoming County, X 4. J. NYSM 12741, cross section based on a specimen from Randall Gully (Loc. 44/4), near Bristol Center, OntarioCounty, X 4. K. NYSM 12742, cross section of a specimen from the Parrish Limestone, Griesa Hill Road (Loc. 46d), Naples, OntarioCounty, X 4. L. NYSM 12740, suture at 17.1 mm diameter, from Randall Gully (Loc. 44/4), near Bristol Center, Ontario County, X 4.M, NYSM 12743, suture at ca. 20 mm diameter based on a specimen from the shale capping the Parrish Limestone, Conklin Gully (Loc.47/2), near Naples, Yates County, X 4. N, NYSM 4067, the holotype (plastotype), reversed suture at ca. 23.5 mm diameter, from the shalecapping the Parrish Limestone from the same locality and horizon, X 4. O, NYSM 12724, reversed suture at 24.8 mm diameter, fromCayuga Creek (Loc. 33/8), Cowlesville, Wyoming County, X 4. P, NYSM 12745, reversed suture at ca. 5.1 mmwhorl height, from the ParrishLimestone, Snyder Gully (Loc. 46c/1), Woodville, Canandaigua Lake, Ontario County, X 4. Q, NYSM 3648, the holotype of Gephyrocerasholzapfeli Clarke, 1898, suture at 16.8 mm diameter recorded as from the lower Portage shales, Eighteenmile Creek, near North Evans, ErieCounty, and probably from the Cashaqua Shale, X 5.


Text-fig. 51. Sutures and cross sections of Probeloceras lutheri (Clarke, 1885) from the Cashaqua Shale of New York State. A. NYSM 12173,cross section at ca. 41 mm diameter, Cazenovia Creek (Loc. 32/8), Spring Brook, Erie County, X 3.2. B. NYSM 12174, cross section, Eigh-teenmile Creek (Loc. 30/8), North Evans, Erie County, X 3.2. C. NYSM 12175, cross section at ca. 28 mm diameter, Little (continued right)


lines of convex type with broad ventral sinus, arched lateralsalient, and indications of umbilical sinus (Text-fig. 49E).

Discussion.–This species is characterized by the well-rounded whorl section throughout ontogeny as illustratedby Glenister (1958: 72, text-fig. 4; UWA 35766) althoughin that specimen, the outermost whorl (at 20.9 cm diameter)has become laterally compressed; it is comparable at similardiameters to the New York material (Table 21). As men-tioned above, one specimen showing growth lines in theAustralian suite shows a slight sinus on the lateral salient(UWA 35774), as does Crickites cordiforme, but the onespecimen from Walnut Creek (Loc. 89a) showing thegrowth lines well (Text-fig. 49E) shows no evidence of this.The specimens assigned to Manticoceras lindneri from

the Cerro Gordo Member of the Lime Creek Formation ofIowa (Baker et al., 1986) lack the distinctively inflated whorlform of the Australian and New York Representatives of thespecies. These Iowa forms are part of the M. regulare faunaof UD I-J (Becker & House, 2000) and could belong toSphaeromanticoceras. TheM. regulare fauna is also found inthe Amana Beds in Iowa (Glenister et al., 2002).

Distribution.–Virgin Hills Formation, Canning Basin,Western Australia. Lower Hanover Shale (West FallsGroup), Walnut Creek (in the vinicity of Loc. 89a/4-5), atSilver Creek (Chautauqua County).

Range.–Frasnian: Crickites Genozone UD I-L. RegionalZone of C. lindneri (I-L1) (Western Australia: Becker et al.,1993; Becker & House, 1997), C. lindneri (24a) (NewYork). Conodont Zone MN 13 (Australia), MN 12-?13(New York).

?Crickites sp. juv.Text-fig. 48H

New material.–NYSM 16589, single small specimenshowing a cross section but neither suture nor growth lines.

Dimensions.–At D = ?4.4 mm, WW = 2.13 mm andWH = 2.3 mm.

Description.–Poorly preserved cross section crushed inearliest whorls. Subinvolute, final whorl section withrounded venter and umbilical slopes with flat flanks con-verging ventrally. Suture and growth lines not seen.

Discussion.–This record, together with ?Archoceras sp.(NYSM 16587), are the youngest Frasnian goniatites knownin New York and thus have considerable importance. Therounded, stout cross section is reminiscent of Crickites

holzapfeli, the highest Frasnian goniatite zone-fossil in Eu-rope, which also survived until the end-Frasnian extinctionevent. A thorough search of this level in Irish Gulf (Loc. 90)is required. That could enable the recognition of a terminalRegional Zone named after the species in New York.

Distribution.–Upper Hanover Shale (West Falls Group),Irish Gulf (Loc. 90/2), near North Boston (Erie County).Bed 90/2 (Text-fig. 16) is a 2.0-2.5 cm-thick crinoidal andgastropod-rich concretionary level, 1.76 m below the Frasn-ian-Famennian boundary and indicated by a concretionsymbol by Over (1997b: fig. 8).

Range.–Frasnian: Crickites Genozone UD I-L. RegionalZone not assigned (24c). Conodont Zone MN 13.

Family BELOCERATIDAE Hyatt, 1884Diagnosis.–Gephuroceratoideans in which sutural ele-

ments become sigmoidal and usually angular with progres-sive increase to seven adventitious and eight or more ventralumbilical lobes with many matched on dorsal suture. Rarelywith one ceratitic lobe. Shell laterally compressed and even-tually discoidal but with characteristic ventral band com-monly rather tabular. Growth lines biconvex.

Discussion.–The relations of this highly distinctive familyhave now been clarified by the New York faunas, especiallythe ancestral Probeloceras (see Kirchgasser, 1975), and doc-umentation of the biostratigraphy of more complex formsfrom the Canning Basin (Becker et al., 1993). This work hasshown that the compilation of Yatskov (1990), which didnot have the advantage of the revision of non-Russian ma-terial nor a stratigraphy, is largely illusory. Nevertheless, weaccept his new genus Naplesites Yatskov, 1990, as a conven-ient name for the intermediate group between Probelocerasand Mesobeloceras Glenister, 1958, and hence to the mostmorphologically complex groups, BelocerasHyatt, 1884, andCeratobeloceras House & Kirchgasser in House et al., 1985.Essential in the simplification has been the removal of eobe-loceratids thought by Yatskov to be contemporary butshown to be late Givetian from work in Morocco (Becker &House, 2000a).There remains the question of the origins of the lineage.

The group seems clearly to have arisen from the Acanthocly-meniidae, and a case can be made that Acanthoclymenia ge-nundewa is the earliest, but the only known material is verysmall, and although some material of that taxon develops asigmoidal ventrad face to the main lateral saddle (Kirch-gasser, 1975: text-fig. 9E), here the definition is based on

(Text-fig. 51 continued) Tonowanda Creek (Loc. 35/8), Linden, Genesee County, X 4. D. NYSM 12177, suture at ca. 11 mm diameterand ca. 4.6 whorl height, Cazenovia Creek (Loc. 32/7), Spring Brook, Erie County, X 10. E. NYSM 12176, reversed suture of a flattenedspecimen at ca. 11 mm whorl height, from the shale on top of the Parrish Limestone, from a gully northeast of Lee Cemetery (Loc.47c/1), Yates County, near Naples, Ontario County, X 6. Scales = 5 mm.


the entry of the larger species, Probeloceras lutheri.Distribution.–North America, Europe, North Africa,

Asia, and Western Australia.Range.–Frasnian: Probeloceras to Crickites Genozones

UD I-E-L (accepting P. lutheri as the base of the range).

Genus PROBELOCERAS Clarke, 1898Type species.–Goniatites lutheri Clarke (1885: 50), by

monotypy.Diagnosis.–Beloceratids with no subdivision of ventral

lobe beyond that found in acanthoclymeniids but with sig-

moidal main lateral saddle, pointed lateral lobe (L), and sub-division of the dorsal suture giving a U1 lobe in adult. Shellform laterally compressed with flattened venter. Growthlines with strong forward projection across flanks.

Distribution.–North America and Western Australia.Range.–Frasnian: ProbelocerasGenozone UD I-E-F. Con-

odont Zones MN 5-6.

Probeloceras lutheri (Clarke, 1885)Pl. 24, Figs 5-9, Pl. 25, Fig. 1; Text-figs 35C, 50-51; Table 22

Goniatites lutheri Clarke, 1885: 50, pl. 2, fig. 8.

Text-fig. 52. Sutures of Naplesites from New York State. A-E. N. iynx (Clarke, 1898), both probably from the lower Rhinestreet Shale atNaples, Ontario County. A-D. Sutural development probably based on NYSM 3583, as illustrated by Clarke (1899: 104): A, at 1.25 whorls;B, at 2.25 whorls; C, at 3.5 whorls; D at 3.75 whorls, magnification not stated. E. NYSM 3586, reversed suture at 22 mm diameter, X 6.F-H.N. naplesense (Clarke, 1898), both probably from the lower Rhinestreet Shale at Naples, Ontario County. F. NYSM 4072, the holotypefigured by Clarke (1899: 105, pl. 7, fig. 18), reversed suture at 33 mm diameter, X 6. G. The same, reversed suture, at 30 mm diameter, X6. H. NYSM 4066, a specimen from among Clarke's syntypes of Probeloceras lutheri (Clarke, 1885), reversed suture at 28 mm diameter, X4.5.


Gephyroceras holzapfeli Clarke, 1898, 1899a, b: 87, pl. 7, fig. 17;text-fig. 65.

Probeloceras lutheri. Clarke, 1899a, b (pars): 90-102, pl. 7, figs 6,8-10 [not figs 1-5 = Prochorites alveolatus (Glenister, 1958),fig. 7 = Naplesites naplesense (Clarke, 1898)], text-figs 68-69,72, 75-76 (not 73-74, 77 = P. alveolatus); Miller, 1938 (pars):63-70, pl. 12, figs 1, 4, 8-10 (not figs 2-3, 5-7 = P. alveolatus),text-figs 10A-B, H, J (not text-figs 10C, E, ?F, G, I = P. alveo-latus); Kirchgasser, 1975: 73-77, pl. 1, figs 1-5, 13-14, pl. 2,figs 7-21, pl. 3, fig. 2, text-figs 4-7, table 1; House & Kirch-gasser, 1993: fig. 4A; Becker, House & Kirchgasser, 1993: figs4C-D.

Type specimen.–Holotype, USNM 14993; NYSM 4067(plastotype), figured here Pl. 24, Figs 7-8, from shale cap-ping Parrish Limestone, Cashaqua Shale, Conklin (Parrish)Gully (Loc. 47/2), near Naples (Ontario County; Kirch-gasser, 1975: 73).

Diagnosis.–Conch relatively large (to D = 60 mm),highly evolute (subserpenticonic), widely umbilicate.Whorlhighly compressed (discoidal) with steep umbilical walls,subparallel, flattened flanks, weak ventrolateral furrows, andflat (tabular) venter that becomes rounded in outermostwhorls. Suture (E1-Em-E1, L, U1, I) distinguished by highlyasymmetrical, angular to acute lobes and saddles in chevronpattern.

Dimensions.–See Table 22.Description.–Highly evolute and widely umbilicate

through all growth stages. Whorls increasingly compressedbeyond 2-3 mm diameter with flanks subparallel at nearlyright angles to steep umbilical walls (Text-figs 35C, 50).Weak ventrolateral furrows developing with whorl compres-sion to latest stages. Venter distinctly flat or tabular by 1.5mm diameter, becoming rounded beyond approximately 22mm diameter.Suture across venter following pattern of Manticoceras

with ventrolateral lobes (E1) arising from bottom rather thansides of ventral lobe (Text-fig. 50A; Kirchgasser, 1975: fig.4A). Sutural ontogeny marked by increasing angularity oflobes and saddles as characteristic chevron pattern develops.Ventral sides of lateral saddle becoming markedly crookedwith distinct reflection toward umbilicus (Text-fig. 50). Inlatest stages, ventrolateral and lateral lobes becoming acuteand lateral saddle nearly so.Growth lines of fine raised lirae with weak salient across

umbilical shoulder, shallow broad sinus on lower third offlank, strong, rounded salient on ventrolateral furrow, anddeep, tongue-shaped ventral sinus. Distinctive feature is pairof low, thread-like, revolving keels (“hyponomic tire” ofClarke, 1899a, b) inside the ventral margin (Kirchgasser,1975: Pl. 2, Fig. 19). Ventral wrinkle layer consisting of uni-form, widely spaced parallel lines that sweep gently forward

across flanks and cross venter at right angles to revolvingkeels.

Discussion.–The strongly asymmetrical and angular lobesand saddles of Probeloceras lutheri serve to distinguish Probe-loceras from Acanthoclymenia and Prochorites. Prochorites isdistinguished by its concave venter.

Distribution.–Lower Cashaqua Shale (Sonyea Group),widespread across outcrop belt between Erie and Steubencounties. Unconfirmed records in Brallier Shale or equiva-lents in Pennsylvania, West Virginia, and Virginia, andWoodmont Shale or equivalents inWest Virginia andMary-land, and Millboro Shale of Virginia (Kirchgasser, 1975;House 1978). Gogo Formation, Canning Basin, WesternAustralia (Becker et al., 1993, Becker & House, 1997).

Range.–Frasnian: Probeloceras to Prochorites GenozonesUD I-E-F. Regional Zone of Probeloceras lutheri (19) to ?Pro-chorites alveolatus (20) (New York). Regional Zones P. lutherilutheri (E1) to Sphaeromanticoceras affine (F3) (CanningBasin, Western Australia). Conodont Zones MN 5-6.In New York, not known with certainty from upper part

of Cashaqua Shale including the Shurtleff Septarian Hori-zon which yields Prochorites alveolatus.

Text-fig. 53. Sutures and growth lines of Tornoceras, from the Ge-nundewa Limestone and Angola Shale, and Crassotornoceras, fromthe Angola Shale in New York. A-B. T. uniangulare compressumClarke, 1897, based on a specimen from the Genundewa Lime-stone, Bethany Center (Loc. 24), Genesee County. A. NYSM12055, suture at ca. 22 mm diameter (estimated), X 1.3. B. NYSM12056, growth line at ca. 28 mm diameter based on a specimenfrom the same locality and horizon, X 1.3. C-D. T. cf. typum (G.& F. Sandberger, 1851), based on a specimen from the AngolaShale in Cazenovia Creek (Loc. 75/6), Erie County. C. NYSM12050, growth line at whorl height of 9.9 mm, X 2.7. D. Sutureat a whorl height of 8.3 mm, X 2.7. E-F. C. aff. crassum (Matern,1931), NYSM 12014, based on a specimen from the Point BreezeGoniatite Bed, Angola Shale, Johnson Creek (Loc. 81/6),Wyoming County, X 3. E. Growth line. F. Course of constriction.


Text-fig. 54. Sutures of Tornoceras spp. and sspp. from the Hamilton Group, Tully Limestone, and Genesee and Sonyea groups in New Yorkand the Squaw Bay Limestone in Michigan, and Phoenixites concentricus (House, 1965), from the Gowanda Shale in New York. A-C. T. uni-angulare uniangulare (Conrad, 1842). A. NYSM/CU 40115, suture at whorl height of 15.2 mm of a specimen from the Leicester ("Tully")Pyrite near Leicester, Livingston County, X 2.5. B. AMNH 5476(ii), suture at 20 mm diameter based on one of Conrad's cotypes from thesame locality and horizon, X 2.5. C. AMNH 5476(i), suture at 12 mm diameter based on one of Conrad's cotypes from the same localityand horizon, X 3.5. D. T. cf. uniangulare, USNM 137721, suture at whorl height of 16.5 mm based on a specimen from the lowerWindomShale (Hamilton Group) at a quarry 1 mi (1.6 km) southeast of Lebanon Center, near Hamilton, Madison County, X 2. E. T. cf. uniangularetoward Epitornoceras, NYSM 12049, suture of a specimen from a level in the Geneseo Shale, 2.4 m (11 ft) above the Leicester Pyrite (Loc.18/2), Beards Creek, Leicester, Livingston County, X 2. F. T. u. compressum Clarke, 1897, NYSM/D 1444, suture at whorl height of 15.3mm based on a specimen from the Genundewa Limestone at Bethany Center (Loc. 24), Genesee County, X 3. G. T. u. obesum Clarke, 1897,NYSM 4093, reversed suture, based on a specimen from the "Naples Group" (probably upper Cashaqua Shale) at Mount Morris, LivingstonCounty, figured by Miller (1938: pl. 31, figs 5-6), X 2. H-J. T. arcuatumHouse, 1965, sutures of specimens from the Squaw Bay Limestoneat Partridge Point, Alpena County, Michigan. H. USNM 96543a, reversed suture at 38.7 mm diameter, X 1.5. I. MMP 47509, suture at27 mm diameter, X 1.2. J. NYSM/D 1430, suture at 20 mm diameter, X 2.5. K. T. cf. arcuatum, USNM 96551a, reversed suture at estimateddiameter of 41 mm, based on a specimen from the Platyceras Bed of the Tully Limestone, West Brook bed or member, (continued right)


Text-fig.55.Graphsshowingtheontogenyofsubspeciesof

Tornoceras

uniangulare(Conrad,1842)from

NewYork;seeTables23-25forbiometricdata.A.T

.u.u

nian

gulare,data

basedonmateriallistedhereinandbyHouse(1965),allfrom

theLeicester("Tully")Pyrite,Leicester,LivingstoneCounty,andCanandaiguaLake.B.T

.u.com

pressumClarke,

1897,databasedonmateriallistedhereinandbyHouse(1965),allfromtheGenundewaLimestone;allthematerialabove12mmdiameter,andmuchofthatbelow,isfromBethany

Center(Loc.24),GeneseeCounty.C.T

.u.obesumClarke,1897,databasedonmateriallistedhereinandbyHouse(1965),allfrom

theCashaquaShaleandmostprobablyfrom

theShurtleffSeptarianHorizonintheHoneoyeLakearea.

(Text-fig.5

4continued)fimbriatazone(Cooper&Williams,1935),0.5mi(0.8mm)northwestofGeorgetown,MadisonCounty,X1.L.

Tornoceras

arcuatum,NYSM

12024,

sutureatca.24.5mmdiameterbasedonaspecimenfrom

theCrosbySandstoneinSunsetPointGully(Py-16/1),EastBranch,KeukaLake,YatesCounty,X1.5.M-O.P.

concentricus,suturesofspecimensfromtheCorell'sPointGoniatiteBedoftheGowandaShaleatCorell'sPoint(Loc.97),Brockton,ChautauquaCounty.M.USNM137667,re-

versedsutureat28mmdiameter,X2.5.N.NYSM

11963,sutureat20mmdiameter,acompositedrawingfromadjacentsepta,X2.5.O.NYSM

11964,sutureat12.9mmdiameter,

X3.P.T.sp.USNM96551D,sutureatestimatedwhorlheightof29mmbasedonaspecimenfromthePlatycerasBedoftheTullyLimestone,WestBrookbedormember,fimbriata

zone(Cooper&Williams,1935),0.5mi(0.8km)northwestofGeorgetown,MadisonCounty,X1.12.


In the Canning Basin,Western Australia, Probeloceraslutheri precedes Prochorites alveolatus as in New York, butunlike in New York, it ranges upward and overlaps P. alveo-latus in the form of P. aff. lutheri (with subacute venter) (F1),P. lutheri n. ssp. (in a sequence linking P. lutheri withNaple-sites showing the gradual entry of an E2 lobe (F2) and P. cf.lutheri associated with Sphaeromaticoceras (Becker et al.,1993; Becker & House, 1997: 141; 2000: 128).

Genus NAPLESITES Yatskov, 1990Type species.–Probeloceras? naplesense Clarke (1898,

1899a, b), by original designation (Yatskov, 1990: 43).Diagnosis.–Beloceratids with single subdivision of ventral

saddle to form E2 lobe, and single subdivision low on flanksto form U2, and perhaps incipient E3 with up to four lobesbetween L lobe and seam; all lobes acute soon after develop-ment; saddles subangular to acute. Intermediate in prolifer-ation of acute lobes and saddles between Probeloceras andMesobeloceras in a lineage that culminates in BelocerasHyatt,1884.

Included species.–Naplesites anguisellatum (Chao, 1956),N. housei (Montesinos & Henn, 1986), N. iynx (Clarke,1898, 1899a, b), N. cf. inyx (Becker et al., 1993), N. naple-sense (Clarke, 1898, 1899a, b),Naplesites sp. (Bogoslovsky etal., 1982: referenced by Becker & House, 2000: 129).

Distribution.–North America (New York; see Adden-dum), Europe, Asia, and Western Australia.

Range.–Frasnian:MesobelocerasGenozone UD I-G. Sub-division UD I-G1, range of Naplesites before the entry ofMesobeloceras (UD I-G2) (Becker & House, 2000).

Naplesites naplesense (Clarke, 1898)Pl. 25, Fig. 6; Text-figs 52F-H

Anabeloceras pseustes Clarke, 1897: 53, nom. nud. et dubium.Probeloceras? Naplesense Clarke, 1898, 1899a, b: 105, text-fig. 78,

pl. 7, fig. 18.Neomanticoceras naplesense. Schindewolf, 1936: 690; Miller, 1938:

135, fig. 28B, pl. 12, fig. 11); House, 1962: 258.Mesobeloceras naplesense. House & Kirchgasser, 1993: 276, 278,

fig. 5J.

Type material.–Yatskov (1990) based his interpretationon the holotype (NYSM 4072; formerly 12481/1), then theonly known specimen figured by Clarke (1899a, b: text-fig.78, pl. 7, fig. 18) and Miller (1938: pl. 12, fig. 11, copiedfrom Clarke). Reexamination of the holotype (by MRH in1959 and subsequently) shows that the suture on the lowerflanks was incorrectly interpreted and that the suture thereactually shows a sharp L lobe and angular U2 probably ven-trad of the seam (revised here; Text-fig. 52G). Thus the as-sigment of Miller to Neomanticoceras is incorrect as is thediagnosis of Yatskov (1990).

New material.–The suture of a better specimen found byWTK among the syntypes of Probeloceras lutheri, which issimilar (NYSM 4066, formerly 12480/6), is illustrated here(Text-fig. 52H).

Dimensions.–Holotype: D = 61 mm; WH = 23.3 mm;UW = 19 mm.

Description.–The holotype is here illustrated photo-graphically for the first time (Pl. 25, Fig. 6). The specimenis crushed flat but consists of a thin, crushed pyritic mold ofthe earlier whorls. The suture, illustrated by Clarke (1898,1899a, b: 105, text-fig. 78) and repeated by Miller (1938:text-fig. 28B), shows only a broadly rounded lateral saddle(L). It was noted by MRH (in 1959) that there was evidenceof two angular lobes before the seam (Text-fig. 52G), andthis is confirmed in the new specimen (NYSM 4066; Text-fig. 52H).

Discussion.–Our reinterpretation of the lateral suture ren-ders the assignment to Neomanticoceras of Schindewolf(1936) andMiller (1938) untenable. A topotype ofN. para-doxum, the type species of Neomanticoceras, fromBüdesheim, Germany, is figured here (Pl. 7, Figs 2-3) to il-lustrate differences. These New York specimens thus occupy

Table 23. Biometric data for Tornoceras uniangulare uniangulare (Conrad, 1842) from the Leicester Pyrite. See Text-fig. 55A for graph.


AMNH 5476 ca. 19 ca. 8.5 11.0 ca. 0 0.77 —(holotype)

NYSM 16539 ca. 30.5 10.0 18.0 ca. 0 0.56 —CU 40115 (iii) 21.5 8.5 11.6 — 0.73 —AMNH 5476 (ii) ca. 20.5 ca. 9.5 — 0 — —NYSM 16540 18.6 7.7 11.6 — 0.66 —CU 40115 (ii) 10.1 4.7 6.4 — 0.73 —NYSM D 1345 7.2 3.8 ca. 4.0 — 0.95 —NYSM 16540 4.4 2.5 2.3 — 1.08 —NYSM 5662 3.9 2.3 1.93 — 1.19 —NYSM 5661 1.36 1.04 0.49 — 2.12 —



an intermediate position between Probeloceras (but with anA2 and single ventral U lobe) and Naplesites iynx (with anacute A2 lobe and two ventral acute U lobes). AlthoughYatskov (1990) intended his genus Naplesites to apply to aquite different form, we have now (with R. T. Becker) doc-umented a succession inWestern Australia through increas-ingly complex sutures to Mesobeloceras (with acute A2, A3,and even in some A4 lobes and with at least three ventral Ulobes). Hence Naplesites has a restricted time span and thename is biostratigraphically convenient.

Distribution.–Probably lower Rhinestreet Shale (WestFalls Group), in the area of Naples, Ontario, and Yates coun-ties. Reported from the “soft shales at Naples” and thoughtto be from the lower Rhinestreet Shale. The reference toNaples in Clarke’s (1899c: 85) locality guide gives no helpon the precise locality among sections in Parrish (Conklin)Gully (Loc. 65a), Grimes Gully, or Tannery Creek. All havebeen searched but no new material has been found.

Range.–Frasnian:MesobelocerasGenozone (inferred) UDI-G. Subdivision UD I-G1, range of Naplesites before entryofMesobeloceras (inferred). Regional Zone of N. iynx (21a).Conodont Zone unknown: MN ?6 or ?7.

Naplesites iynx (Clarke, 1898)Pl. 25, Figs 2-5; Text-figs 52A-E

Beloceras iynx Clarke, 1897, nom. nud.; Clarke, 1898, 1899a, b:103-104, text-figs 77-79, pl. 7, figs 11-16.

Probeloceras iynx. Wedekind, 1918: 131.Eobeloceras iynx. Schindewolf, 1936: 690; Miller, 1938: 137-139,

text-fig. 29, pl. 30, figs 12-17.Naplesites clarkei Yatskov, 1990 [based on Clarke 1899a, b: pl. 7,

figs 15 (NYSM 3585) and 16 (NYSM 3586), text-fig. 78(NYSM 3583), and another specimen].

Mesobeloceras iynx. House & Kirchgasser, 1993: 276, 278.

Type material.–Lectotype designated by Yatskov (1990:43) is NYSM 3581 (formerly 12060/1). Other cotypesmentioned below, assigned elsewhere by Yatskov (1990) butregarded here as synonyms of Naplesites iynx, include:NYSM 3582 (Pl. 25, Fig. 2), 3583 (Pl. 25, Fig. 4), and 3584(not illustrated here), the umbilical portion of which is bro-ken off, so the umbilical lobes are not seen, hence the as-signment is not clear. The original syntypes were formerlynumbered NYSM 12060/1-6.

Dimensions.–Lectotype, NYSM 3581: D = 25.4 mm,WW = 11.2 mm, UW = 7.3 mm.

Diagnosis.–Distinguished from Naplesites naplesense byadditional sharp umbilical lobe on low flanks.

Description.–Clarke (1899a, b) and Miller (1938) gavedetailed descriptions of the type material and four of thesespecimens are here illustrated photographically for the first

time. The inner whorls are pyritic and three-dimensionalbut the outer whorls are crushed. The diagram of early su-tural ontogeny (Clarke, 1899a, b: text-fig. 77; Miller, 1938:text-fig. 29) is reproduced here (Text-figs 52A-E); in truththe specimen on which this ontogeny is based (NYSM3583; Pl. 25, Fig. 4) could also represent early stages ofNaplesites naplesense, which would be indistinguishale fromN. iynx in early stages.

Discussion.–A rather similar and possible junior synonymof Naplesites iynx is N. anguisellatum, the type species ofChaoceras Yatskov, 1990, a genus here regarded as a juniorsynonym of Naplesites.The sequence of morphologies between Probeloceras and

early Mesobeloceras in Western Australia, which includeNaplesites cf. iynx and N. housei, indicates that the evolu-tionary stages were short lived (Becker et al., 1993; Becker& House, 1997, 2000). The rarity of N. naplesense and N.iynx suggest that they could have been closely contempo-rary.

Distribution.–Probably lower Rhinestreet Shale (WestFalls Group), in the area of Naples, Ontario. and Yates coun-ties. Reported by Clarke from Portage shales at Naples butno locality or horizon details were given (see Distributionof Naplesites naplesense).

Range.–Frasnian: Mesobeloceras Genozone ((inferred)UD I-G. Subdivision UD I-GI, range of Naplesites beforeentry of Mesobeloceras (inferred). Regional Zone of N. iynx(21a). Conodont Zone unknown: MN ?6, or ?7.

Order GONIATITINA Hyatt, 1884Superfamily TORNOCERATOIDEA Arthaber, 1911

This is the longest ranging superfamily of coiled ammonoidsin the Devonian and, during its range, perhaps the com-monest. The superfamily is characterized by a generally sim-ple suture comprising a ventral lobe (E), an adult lateral lobeon the flanks, which develops adventitiously during on-togeny (A), a lobe centered on the seam (L), and a dorsallobe (D). Later forms can be more complex. Growth linesare normally biconvex. Tornoceratids first appear in the lateEifelian and continue to the latest Famennian, the lastrecords being from theWocklumeria Stufe when extinctionis associated with the Hangenberg Event. The group is wide-spread and cosmopolitan and is known from all continentsexcept for polar regions.Three families are included: Tornoceratidae, Post-

tornoceratidae, and Pseudoclymeniidae. Only the first isknown in New York where it has a long record from bedsimmediately below the Cherry Valley Limestone (HamiltonGroup) to the Gowanda Shale (Canadaway Group).

Family TORNOCERATIDAE Arthaber, 1911

Diagnosis.–Tornoceratids with imperforate, ornamentedammonitella; umbilici and umbilicus open or closed withshell form rotund and rounded to compressed and platy-conic or oxyconic. Adult shell usually smooth, rarely withfalcate lirae, prominent ribs, nodes, or constrictions. Sutureas described for superfamily but with simplification of lobesin some (Tornia and Kirsoceras) and additional elements inothers, a dorsal umbilical lobe (U), for example (in Loboto-rnoceras and Gundolficeras).Protoconch and ammonitella, where known, orna-

mented with radial lirae; growth lines primarily biconvexafter nepionic constriction. Dorsal wrinkle layer spirallingbackward from umbilicus to become almost rectiradiate overventer.

Discussion.–Several distinct groups are included withinthe family, which is thought to be derived from an unspec-ified Anarcestidae source. Only the family Tornoceratidae isof concern here because the other families, Posttornocerati-dae (developing additional ventral adventitious lobes) andPseudoclymeniidae (with clymeniid-like lateral lobes), arenot represented in North America.

Subfamily PARODICERATINAE Petter, 1959The Parodiceratinae forms the earliest group and comprisesParodiceras, with only an incipient lateral lobe (A), as theearliest member, andWedekindella and Trevoneites. The lasttwo, reviewed by Becker & House (1994b), are only tenta-tively included here; both have well-developed lateral (A)lobes butWedekindella is a constricted form and Trevoneiteshas an open umbilicus. Only Parodiceras occurs in New Yorkin the Union Springs Shale (Werneroceras Bed) and CherryValley Limestone (House, 1981b, 1978) of the HamiltonGroup, well below the levels dealt with in this monograph.

Subfamily TORNOCERATINAE Arthaber, 1911Tribe TORNOCERATINI Arthaber, 1911

Diagnosis.–Tornoceratidae with smooth shells, roundedventers, no ventrolateral furrows or tabular venters, and gen-erally with umbilici.

Discussion.–This group includes: TornocerasHyatt, 1984,the long-ranging nominal genus; Linguatornoceras House,1965, a group with lingulate rather than asymmetric lateral(A) lobes; Epitornoceras Frech, 1902a, a late Givetian andearly Frasnian oxycone; Crassotornoceras House & Price,1985, a late Frasnian constricted form; and LobotornocerasSchindewolf, 1936, which adds a small saddle on the seam.All of these occur in New York. Other genera are: Oxy-tornoceras Becker, 1993a, a Famennian homeomorph of Epi-tornoceras;Domanikoceras Becker & House, 1993, a strangeearly Frasnian form with convex growth lines from theTiman; and Semicheiloceras Becker, 1993a. These last threegenera are not known in North America.

Distribution.–Worldwide. Marcellus Formation (Hamil-ton Group) to Gowanda Shale (Canadaway Group) in NewYork.

Range.–Givetian MD II-A to Famennian UD VI-D.

Genus TORNOCERAS Hyatt, 1884Type species.–Goniatites uniangulare Conrad, 1842.Diagnosis.–Involute, well-rounded, or subglobular to

subplatyclonic shell with closed or nearly closed umbilicus;without marked ventrolateral furrows. Adult suture withEALD lobes only. Adult growth lines commonly biconvexbut biconvexity lost toward aperture in some species.

Discussion.–Revision of the species of Tornoceras is stillin progress. Several dozen names are taxonomically availablefor this genus and these will only be referred to in the follow-ing text if they are relevant to New York material.

Table 24. Biometric data for Tornoceras uniangulare compressum Clarke, 1897, from the Genundewa Limestone. See Text-fig. 55B forgraph.


NYSM 4098 16.0 ca. 5.5 — — — —(holotype)

NYSM D. 1447 36.0 15.5 20.0 ca. 0 0.78 —NYSM 16547 34.8 14.9 18.3 ca. 0 0.81 —NYSM 16543 30.4 13.6 16.5 ca. 0 0.82 —NYSM 12065 29.2 13.7 15.3 ca. 0 0.90 —NYSM 16542 27.8 11.9 16.7 ca. 0 0.71 —NYSM 16544 22.8 11.2 14.2 ca. 0 0.79 —NYSM 16546 17.7 7.9 10.9 ca. 0 0.73 —NYSM 16548 16.0 7.9 8.8 ca. 0 0.90 —NYSM 16545 11.5 6.0 6.4 ca. 0 0.94 —NYSM 16549 9.7 5.0 5.4 ca. 0 0.93 —CU 101/ix 3.0 1.8 1.6 0 1.13 —



Forms with small lingulate lateral lobes have been sepa-rated as the subgenus Linguatornoceras, elevated herein togenus, as discussed below. The North American (House,1965), Russian (Bogoslovsky, 1971), and German (Becker,1993a, 1995) species of Tornoceras have been subject to re-vision; the varied forms from elsewhere are mostly poorlyknown. A wide range of distinct sutural, growth line, andshell form types are included in the European faunas andthere is no doubt that they will contribute to the discrimi-nation when restudied. Because the New York forms havebeen relatively recently described (House, 1965), this ac-count will be limited to new records and new statistical in-formation.

Distribution.–Extraordinarily wide distribution. Knownon all continents except Antarctica. In New York from Chit-tenango Shale of Marcellus Formation (Hamilton Group)to Corell’s Point Goniatite Bed, Gowanda Shale (Canad-away Group).

Range.–International range: Givetian MD II-A to Fa-mennian UD II-D.

Tornoceras uniangulare uniangulare (Conrad, 1842)Pl. 26, Figs 1-4, 6; Text-figs 54A-C, 55A; Table 23

Goniatites uniangularis Conrad, 1842: 268, pl. 16, fig. 4.Goniatites astarte Clarke, 1885: 29, pl. 2, figs 9-10.Tornoceras uniangularemut. astarte. Loomis, 1903: 916-917, 919,

pl. 5, figs 1-2.Tornoceras (Tornoceras) uniangulare.Miller, 1938 (pars): 157-166,

pl. 31, figs 8-9, pl. 32, fig. 9, pl. 33, figs 5-6, pl. 35, figs 5-6.Tornoceras (Tornoceras) uniangulare uniangulare. House, 1965:

104-106, text-figs 9A-H, 17E, pl. 7, figs 58-59, 61-63, 65-67.

Type material.–The cotypes of Tornoceras uniangulare areAMNH 5476 (two specimens). The one figured by House(1965: pl. 7, figs 65-67, and herein, Pl. 26, Figs 1-2), asAMNH 5476/1, is designated lectotype if it is not the holo-type (it is the only originally figured specimen). The originalspecimens of Goniatites astarte are NYSM 5661 and 5662.All of these specimens are from the Tully (now Leicester)Pyrite.

New Material.–Although additional material of this va-riety has been collected in the present survey, there is noth-ing to add to the general description (House, 1965).Additional statistical information is added to former data inthe graphs (Text-fig. 55A). Sutures illustrative of the sub-species are given inText-figs 54A-C. The material of Loomis(1903; NYSM 5661 and 5662) is from Canandaigua Lake.Other material was listed by House (1965). Our collectionsinclude NYSM 3180/1-3 from the Leicester Pyrite atTaunton Gully (Loc. 17), Leicester, and 3181, 3187/1-3,and 3182 from the same level in Fall Brook (Loc. 19), Gene-

seo. This subspecies is only known with certainty from theLeicester Pyrite.

Dimensions.–See House (1965), Table 23, and Text-fig.52A.

Description.–A description of this subspecies was givenby House (1965).

Discussion.–See House (1965) for detailed discussion ofontogeny of this subspecies.This group represents perhaps the commonest Devonian

goniatite. Comment is necessary, however, on the relativeclaims of Tornoceras uniangulare and Ammonites simplex (vonBuch, 1832: 42, pl. 2, fig. 8), the name commonly used forthis group in Europe. A general comparison between thetwo species has long been claimed, and von Buch’s name forthe European form has a priority of ten years over that fromNew York. Von Buch’s specimen, however, has never beenfigured satisfactorily, and the small sutural thumbnail sketchgiven by him is quite inadequate to form a diagnosis. Fur-thermore, von Buch recorded his specimen “am Rammels-berge bei Goslar gefunden,” in which case the specimen ispossibly a Foordites. Beyrich (1884: 212), however, assertedthat this was an error, and that the specimen was fromBüdesheim, although the grounds for this statement are notclear. If Beyrich was correct, then the specimen probably isa tornoceratid. But in the absence of a clear designation anddescription, it seems that the species is critically undeter-mined. It is true that Beyrich argued that the specimen fig-ured by the Sandberger brothers (1851: pl. 10, fig. 14) asGoniatites retrorsus var. typus, was the same species, and ithas long been been interpreted as that in Germany, but therehas been no formal designation of a neotype for A. simplex.It is notable that the species and subspecies referred to

Tornoceras uniangulare by House (1965), in which they showadult growth lines, give indication of a loss of biconvexity to-ward the aperture; these are all Middle Devonian or earlyUpper Devonian specimens and range to the GenundewaLimestone. On the other hand, T. simplex and T. typum arereported to have biconvex growth lines, the feature of earlystages of T. uniangulare. It could be that the later Tornoceras,from the Büdesheim level, differ in this respect from T. uni-angulare, but in the absence of modern work on EuropeanFrasnian tornoceratids, this view is largely speculative.

Distribution.–Leicester Pyrite (Genesee Group), Leices-ter (Livingston County). The two specimens of Hall(AMNH 5476) were from the Leicester (Tully Pyrite) nearLeicester (Livingston County), but which of the several gul-lies there (Loc. 17, 17a, or 18) is not recorded. The materialof Loomis (NYSM 5661 and 5662) is from CanandaiguaLake. Other material is as reported by House (1965). Fol-lowing the work of Huddle (1974, 1981) and Baird & Brett(1986a), it is recognized that the Leicester Pyrite is a remanié


level that youngs progressively to the west and includes ma-terial derived from the upper Hamilton Group.

Range.–Probably Givetian MD II, now found derived inGivetian MD III.

Tornoceras uniangulare (Conrad, 1842) GroupThere is a range of poorly preserved material that does notallow the rigorous analysis applied to the various subspecies,and that is referred to the Tornoceras uniangulare group invarious forms of open nomenclature as listed below. Theprecise determination of this material is impossible becausemuch of it is crushed and shows no sutures.

Tornoceras aff. uniangulare (Conrad, 1842)Pl. 27, Figs 1-6

Material assigned to Tornoceras (T.) aff. uniangulare was de-scribed and figured by House (1965: pl. 10, figs 92-93, 97-100) from the upper meter or so of the Windom Shale(Moscow Formation, Hamilton Group) in Grove’s CreekQuarry (Seneca County; Loc. 4; NYSM/CUPM 42402,NYSM 12641-12643). There is a larger specimen of T. cf.uniangulare from the Windom Shale in a quarry 1 mi (1.6km) southeast of Lebanon Center, near Hamilton (MadisonCounty; USNM 137721) that might belong here.

Tornoceras cf. uniangulare (Conrad, 1842)Text-fig. 54D

The following material is referred to Tornoceras (T.) cf. uni-angulare: USNM 137721 from theWindom Shale (MoscowFormation, Hamilton Group) and near the top of the unitat Portland Point Quarry on the eastern side of CayugaLake, a large phragmocone and body chamber collected byW. A. Oliver, Jr. (NYSM 12031). From the Geneseo Shale(Genesee Group) on Beards Creek, Leicester (LivingstonCounty; Loc. 18/1, 3108), over 30 poorly preservedTornoceras from 7.6 cm (3 in) below the first 2.5 cm (1 in)black shale. From the Penn Yan Shale, specimens conform-ing with the species occur, including in the Lodi Limestoneat Beards Creek (Loc. 18/4, 3153), a nodule bed 9.75 m (32ft) above the base of the Geneseo Shale. Other materialcomes from a creek north of Abbey Gulf (Loc. 21), 2 mi(3.2 km) northeast of Honeoye (Ontario County), from alevel equivalent to Loc. 21/2 (Linden Horizon), 6.17 m(20.25 ft) below the base of the Genundewa Limestone.Also conforming to the species is the following materialfrom the West River Shale in Livingston County: BeardsCreek (Loc 18a/3), Leicester, 3128/1; Fall Brook (Loc.19a/2), Geneseo, 3117/2; and from higher in the same sec-tion (Loc. 19a/4), 3118/2.

Tornoceras cf. uniangulare (Conrad, 1842)toward Epitornoceras

Text-fig. 54E

There is a single specimen (NYSM 12049) referred herefrom the Geneseo Shale, 2.4 m (11 ft) above the LeicesterPyrite on Beards Creek, Leicester (Livingston County; Loc.18/2).

Tornoceras uniangulare compressum Clarke, 1897Pl. 28, Figs 1-5; Text-figs 53A-B, 54F, 55B; Table 24

Tornoceras uniangulare var. compressum Clarke, 1897: 54; 1898:54; 1899a, b: 116-118, pl. 8, fig. 18 (pars); 1904: 358, 361,371.

Tornoceras (Tornoceras) uniangulare (pars). Miller, 1938: 158, pl.32, fig. 8.

Tornoceras (Tornoceras) uniangulare compressum.House, 1965: 109-110, text-figs 11A-E, 18A.

Type material.–Clarke’s holotype (NYSM 4098) has beendescribed by House (1965).

New material.–Eighty specimens from the GenundewaLimestone at Bethany Center (Genesee County; Loc. 24),formed the basis of the description published earlier (House,1965). There are 47 specimens in the J. W. Wells collection(his loc. 101, NYSM/CU) and 33 specimens (includingNYSM 12634-12640) collected by MRH and used in the1965 study. In addition, there are 20 specimens (includingNYSM 12125-12127) more recently collected. From thislocality, approximately 30 additional specimens are nowavailable. All are solid specimens, occasionally testate, pre-served in coarsely crystalline styliolinid limestone and manyshow sutures and growth lines.

Dimensions.–See House (1965) and Table 24.Discussion.–The holotype shows markedly biconvex

growth lines. The larger Bethany Center specimens differ inshowing a subdued ventrolateral salient and growth linesthat are convex and almost rectilinear across the flanks (Text-figs 53A-B). The accompanying graph (Text-fig. 55B) plotsdimensions of the material available and adds further detailto that previously published. The criteria for the distinctionof this subspecies remains as stated previously: “the consis-tently high whorl-width proportion, the tendency to form atabular venter at high diameters, and the marked reductionin the prominence of the ventrolateral salient above 20 mmdiameter” (House, 1965: 110).

Distribution.–Genundewa Limestone (Genesee Group;Genesee, Livingston, Ontario, and Yates counties).The holotype (NYSM 4098) is labeled “Genundewa

Limestone, Canandaigua Lake.” The new material is mainlyfrom the embankment spoil extracted from the roadcut at

Bethany Center (Genesee County; Loc. 24). So far, the sub-species is only recognized in the Genundewa Limestone. Noattempt was made to search and collect in detail at other lo-calities. Specimens probably belonging to this subspecieswere collected by one of us (WTK) at the type locality ofthe Genundewa at Genundewa Point (Loc. 23b),Canandaigua Lake (Yates County; Loc. 23b/4, 3916; Loc.23b/5, 3915).

Range.–Frasnian: Koenenites Genozone UD I-B. Re-gional Zone of K. styliophilus kilfoylei (16). Conodont ZoneMN 2.

Tornoceras uniangulare obesum Clarke, 1897Pl. 28, Figs 6-10; Text-figs 54G, 55C; Table 25

Tornoceras uniangulare var. obesumClarke, 1897: 54; 1898, 1899a,b: 116, pl. 8, fig. 17.

Tornoceras (Tornoceras) uniangulare (pars). Miller, 1938: 157-166,pl. 32, fig. 7.

Tornoceras (Tornoceras) uniangulare obesum. House, 1965: 110-112, pl. 8, fig. 72, pl. 9, figs 75-79, text-figs 12, 18B.

Type material.–The holotype (NYSM 4099) is a bariticreplacement from the Cashaqua Shale. A description wasgiven by House (1965).

New material.–Two hundred additional specimens fromClarke’s collection (NYSM E 346) of baritic material formedthe basis for the account of this subspecies given earlier: alsoNYSM 11262-11263, 11245, 16564 (D1365), and NYSM4097 and 4093.

Dimensions:–See Table 25 and Text-fig. 55C.Description.–Little addition to the earlier account

(House, 1965) is required. Statistical data are plotted inText-fig. 55C for comparison with the other subspecies. Theregular, sharp fall in the WW/WH ratio has not been seenin other species of Tornoceras. Whereas there are patterns inthe WW/WH ratio that are discriminatory, it should benoted that this material is all baritic shell replacement ma-terial, and hence the measurements refer to testate speci-mens, whereas those for the subspecies uniangulare andcompressum do not. The pattern of sutures is as in T. unian-gulare.

It is material of this subspecies that has provided the bestillustrations available of the wrinkle layer structures inTornoceras (House, 1965: pl. 9, figs 78-79) and larger illus-trations of this are given here: general descriptions have beenpublished by House (1971: 27).

Distribution.–Cashaqua Shale (Sonyea Group; Liv-ingston and Ontario counties).The holotype (NYSM 4099) and also NYSM 11262-

11263, E 346 (approximately 200 specimens) are all bari-tized and said to come from concretions in the CashaquaShale in the Honeoye Lake area. It seems probable that theyare from the Shurtleff Septarian Horizon (Text-fig. 10).Other material is from the type locality of this horizon atShurtleff ’s Gully (Loc. 41/1) (NYSM 16564, = D1365),near Livonia (Livingston County). Specimens more uncer-tainly placed here include NYSM 11245 fromMount Mor-ris (Livingston County), and NYSM 4097 from Naples(Ontario County).

Range.–Frasnian: ProchoritesGenozone UD I-F. RegionalZone of P. alveolatus (20). Conodont Zone MN 6.

Tornoceras arcuatum House, 1965Pl. 26, Figs 7-10, Pl. 27, Fig. 16; Text-figs 54H-J, L

Tornoceras (Tornoceras) uniangulare (pars). Miller, 1938: 165, pl.33, figs. 3-4, pl. 34, figs 1-4; Stumm, 1951: 35.

Tornoceras (Tornoceras) arcuatumHouse, 1965: 106-109, text-figs10, 17D, pl. 8, figs 68-71.

Type material.–The holotype (USNM 96543a) is fromthe Squaw Bay Limestone of Michigan as is all the materialthat formed the basis for the original description.

New York material.–Two specimens (NYSM 12024 and12051), both from the Crosby Sandstone at Keuka Lake(Yates County), are assigned to the species. These are well-preserved phragmocones but without evidence of shell struc-ture.

Discussion.–The two New York specimens are illustratedhere (Pl. 26, Figs 7, 10, Text-fig. 54L) so a detailed descrip-tion is not warranted. The distinctively arcuate latero-umbil-ical saddle, which is nearly symmetrical, is diagnostic of thespecies and well shown in the new material.

Table 25. Biometric data for Tornoceras uniangulare obesum Clarke, 1897, from the Cashaqua Shale. See Text-fig. 55 C for graph.


NYSM 4099 12.1 7.3 — 0 — —(holotype)

NYSM 4093 45.0 16.5 26.0- — 0 0.63 —NYSM 11263 17.8 7.7 11.1-5.5 0 0.69 —NYSM E 346.ii 7.8 4.7 4.8-2.3 0 0.98 —NYSM E 346.i 4.9 3.0 2.8-1.3 0.3 1.07 0.06



House (1965: 108) described a specimen (USNM96551a) from the Platyceras Bed of the Tully Limestone,West Brook bed or member, fimbriata zone (Cooper &Williams, 1935: 804-805), 0.5 mi (0.8 km) northwest ofGeorgetown (Madison County), as having a similar sutureform to this species. There are slight differences in Tornocerascf. arcuatum, so far as the poor state of the specimen allowscomparison, in the different depth of the ventral lobe, andthe lesser symmetry of the latero-umbilical saddle (Text-fig.54K). Nevertheless, the T. arcuatum tornoceratid type couldwell be a long ranging one, starting in the late GivetianPharciceras Stufe and ancestral to the Epitornoceras group, assuggested earlier (House, 1965: 82), although the Michiganand Keuka Lake material is obviously later in age.The occurrence of this species at levels with Koenenites in

the late Penn Yan Shale of New York suggests that theCrosby Sandstone level of Torrey et al. (1932) is a close cor-relative of the Squaw Bay Limestone.

Distribution.—Squaw Bay Limesone, southern side ofPartridge Point, Alpena, Michigan. Crosby Sandstone, PennYan Shale (Genesee Group; Yates County). NYSM 12024,gully above Sunset Point (Loc. F, Py-16/1), Keuka Lake andNYSM 12051, Willow Grove Creek, Keuka Lake (YatesCounty; Loc. E, Py-14/1).

Range.–Frasnian: Koenenites Genozone UD I-B. Re-gional Zone of K. styliophilus styliophilus (16). ConodontZone MN 2.

Tornoceras cf. typum (G. & F. Sandberger, 1851)Pl. 27, Fig. 15; Text-figs 53C-D

Goniatites retrorsus typusG. & F. Sandberger, 1851: pl. 10, figs 14,14a-c.

Type material.–The lectotype is the specimen figured bythe Sandberger brothers and in the Wiesbaden Museum.

New York material.–Three specimens from theRhinestreet and Angola Shales (NYSM 12050, 12076, and12077), preserved as internal molds in mudrock and crys-talline calcite and variously preserving part of the bodychamber and phragmocone. Also a number of very poorlypreserved specimens from the Angola Shale that might be-long here but that cannot be dogmatically assigned.

Dimensions.–Lectotype: D = 45 mm, WW = 18.7 mm,WH = 26 mm, Wh = 14.5 mm, UW = 0 mm.

Description.–The Sandbergers’ drawing of the lectotypeis accurate except that the lateral lobe is more asymmetricaland less tongue-shaped than indicated and the ventrad slopeof the latero-umbilical saddle is almost concentric. Growthlines show a salient low on the flanks, so they are biconvexeven at the large diameter of the specimen.

New York material: closed umbilicate with gentle slopefrom umbilicus to very low shoulder; maximum width verylow on flanks, which converge convexly and smoothly towell-rounded venter. Suture showing gently arched latero-umbilical saddle with flat top (Text-fig. 53D). Growth linesbioconvex with subdued salient low on flanks and promi-nent ventrolateral salient. Weak spiral ornament on flanks.

Discussion.–It is only the evidence of the late stage atwhich biconvexity of the growth lines is retained that justi-fies taxonomic separation from Tornoceras uniangulare, butinsufficient material is available to demonstrate this withcomplete satisfaction given that most specimens do notshow growth lines.

Distribution.–Rhinestreet Shale and Angola Shale (WestFalls Formation; Erie, Wyoming, and Livingston counties).Rhinestreet Shale (approximate level of Relyea Creek

Horizon): NYSM 12076, Cazenovia Creek (Erie County;Loc. 54/1); NYSM 12077, Stony Creek,Warsaw (WyomingCounty; Loc. 61/1). Angola Shale (Point Breeze GoniatiteBed): NYSM 12050, Cazenovia Creek (Erie County; Loc.75/6).NYSM 4093 (Pl. 27, Fig. 15), recorded from the

“Naples Formation” at Mount Morris (Livingston County),is referred here with doubt.

Range.–Frasnian: Playfordites to Neomanticoceras Geno-zones UD I-I-J (also possibly earlier: Mesobeloceras UD I-G). Regional Zone of P. cf. tripartitus (22a) toSphaeromanticoceras rhynchostomum (22b) [also possibly ear-lier: Naplesites iynx (21a)]. Conodont Zone MN 11 to ?12,possibly earlier.

Genus LINGUATORNOCERAS House, 1965Type species.–Goniatites retrorsus var. linguaG. & F. Sand-

berger, 1851.Diagnosis.–Species close to Tornoceras with well-rounded

to compressed-rounded form; distiguished by suture withsmall, deep, usually symmetrical, tongue-shaped lateral lobe.

Included species.–Goniatites lingua G. & F. Sandberger(1851: pl. 10, figs 20-21; 1852: 109); Tornoceras (Lingua-tornoceras) aff. linguumHouse (1965: 112, pl. 9, figs 82, 93,text-fig. 13E); T. clausumGlenister (1958: 92, pl. 15, figs 7-9, text-figs 16a, c); T. guestphalicum Frech (1897: pl. 32a,fig. 8, holotype noted by Becker, 1993a: 188, as refigured byHouse & Price, 1985); T. haugi Frech [1902b: 47, pl. 3(2),figs 20a-b]; T. pompeckjiWedekind [1918: 137, nom. nud.,lectotype designated by Becker, 1993a: 189, as Frech, 1902:pl. 4, figs 9b-c (as refigured by House & Price, 1985)].

Discussion.–Bogoslovsky (1971: 52) did not favor sepa-ration of these forms even at the subgeneric level; this has,however, now been generally adopted. House (1965: 91)thought that the group formed a discrete type of tornocer-

atid restricted to the mid-Frasnian to lower Famennian andwas a useful time discriminator, and this has been confirmedby Becker (1993a). There are other tornoceratids with verylarge lingulate lateral lobes that are nearly symmetrical; theseseem too close to the Tornoceras s. s. stock to warrant taxo-nomic separation.

Distribution.–North America (New York), Europe (Eng-land, France, Germany, Poland), Russia (Urals), NorthAfrica, and Western Australia.

Range.–Frasnian: Prochorites Genozone (UD I-F) toParatorleyoceras Genozone (UD II-D).

Linguatornoceras aff. linguum (G. & F. Sandberger, 1851)Pl. 27, Fig. 12, Pl. 33, Figs 1-2, 5-8; Text-fig. 56; Table 26

aff.Goniatites retrorsus var. linguaG. & F. Sandberger, 1851: pl. 10,figs 20-21; 1852: 109.

Tornoceras (Linguatornoceras) aff. linguum. House, 1965: 112, pl.9, figs 82-83, text-fig. 13E.

Type material.–There has been no further description ofthe type material published since it was originally figuredby the Sandberger brothers.

NewYork material:–Four specimens from the RhinestreetShale referred here (NYSM 12017-12020) and three otherswith some doubt (NYSM 12021-12023), all preserved aspartial calcitic internal molds set in gray mudrock. AlsoNYSM 11248 described by House (1965: 112).

Dimensions.–See Table 26.Description.–Shell involute, with closed umbilicus except

in earliest whorls; well-rounded and ovoid, laterally com-pressed with maximum whorl width low on flanks and curv-ing evenly to seam with no overhang. Suture as illustrated inText-figs 56A-B, D. Growth lines biconvex, with low umbil-ico-lateral and ventrolateral salients, becoming more promi-nent in outer whorls (Text-fig. 56C).

Discussion.–These specimens seem identical with thatfigured by Frech [1902a: pl. 3(2), figs 21a-b] fromBüdesheim under the name Tornoceras simplex mut. ovata(Münster, 1832) but, as Frech recognized, Münster’s nameovata refers to a different form altogether. Linguatornoceraslinguum, on the other hand, differs from the New York spec-

imens in having flat, converging sides, rather than an ovoidwhorl section, and in having a substantially deeper lingulatelateral lobe. It shares the latter character with L. clausumfrom Western Australia. There are specimens from theholzapfeli Zone in Devon, England, figured here as L. aff.linguum (Text-figs 56E-F), which are younger in age andwhich have a flatter and broader ventrolateral saddle. Spec-imens of L. linguum from Germany are illustrated here forcomparison (Pl. 27, Figs 13-14).

Distribution.–Rhinestreet Shale (Erie and Wyomingcounties).Rhinestreet Shale (Relyea Creek Horizon or near equiv-

alent): NYSM 12017 and 12021, Cazenovia Creek (ErieCounty; Loc. 54/1); NYSM 12018 and 12022, RelyeaCreek (Wyoming County; Loc. 60/3); NYSM 12019,Varysburg (Wyoming County; Loc. 57/2); NYSM 12020and 12023, Johnson Creek (Wyoming County; Loc. 58/1).Angola Shale (Point Breeze Goniatite Bed): possible rep-

Table 26. Biometric data for Linguatornoceras aff. linguum (G. & F. Sandberger, 1851) from the upper Rhinestreet and lower Angolashales.


NYSM 12020 21.3 11.5 12.8; ca. 7.5 0 0.90 —NYSM 11248 16.6 8.5 9.0; — 0 0.94 —NYSM 12019 8.9 5.4 5.2; — 0 1.04 —NYSM 12021 4.3 2.7 —: — ca. 0 — —

Text-fig. 56. Sutures and a growth line of Linguatornoceras aff. lin-guum (G. & F. Sandberger, 1851) from New York and Devon,England. A-C. Sutures and a growth line based on NYSM 12020from upper Rhinestreet Shale equivalents on Johnson's Creek (Loc.58/1), Wyoming County. A, Suture at ca. 20.2 mm diameter; B,suture at 13.8 mm diameter; C, growth line at ca. 14.8 mm diam-eter, X 1.5. D. NYSM 11248, suture at ca. 13 mm diameter basedon a specimen collected by D. D. Luther at Relyea Creek (Gibson'sGlen) (Loc. 60), Wyoming County, X 2.7. E-F. Specimens fromthe Saltern Cove Goniatite Bed (holzapfeli Zone), Saltern Cove,Devon, England, sutures reversed for comparison. E. SMH 1526,X 3.7. F. BM c18461, X 3.7.


resentatives include NYSM 3230/13, Cazenovia Creek (ErieCounty; Loc. 75/6).NYSM 11248, collected by D. D. Luther in 1897, was

reported from Relyea Creek (Gibson’s Glen) (WyomingCounty; Locs 60 and 82), but whether it was from the upperRhinestreet or lower Angola is not known (House, 1965).Forms closely related to these specimens from New Yorkoccur in Europe in the upper cordatum Zone and holzapfeliZone of former usage, but they have not been adequatelystudied.

Range.–Frasnian: In New York Playfordites GenozoneUD I-I to ?Neomanticoceras Genozone UD I-J. RegionalZone of P. cf. tripartitus (22a) to ?Sphaeromanticoceras rhyn-chostomum (22b). Conodont Zone MN 11-?12.

Genus CRASSOTORNOCERAS House & Price, 1985Type species.–Tornoceras ausavense crassumMatern, 1931a,

by original designation.Diagnosis.–Tornoceratids with globular inner whorls and

very narrow umbilici with periodic constrictions followingcourse of growth lines and numbering approximately fourper whorl, increasing in some specimens in outer whorls toeight. Conch generally small. Sutures typically simpletornoceratid; some with lingulate lateral lobe.

Included species.–Tornoceras ausavense crassum (Matern,1931a: 27, pl. 3, figs. 14a-b; type species); Crassotornocerasannissi House & Price (1985: 168-169, pl. 16, figs 12-13);T. belgicum (Matern, 1931b: 9, text-fig. 2, refigured byHouse & Price, 1985: text-fig. 3c, pl. 16, figs 1-3); ?C. iso-latum Becker (1993a: 186, text-fig. 68e, pl. 3, figs 11-12);C. nitidum Becker (1993a: 185, text-fig. 68f, pl. 3, figs 15-17).

Remarks.–This distinctive group of small, constrictedtornoceratids seems to characterize the upper part of theFrasnian and the lowest Famennian. There are affinitiesshown by Crassotornoceras both with Linguatornoceras, Lo-botornoceras, and constricted species of Aulatornoceras, butthe exact relationships are obscure.

Distribution.–North America and Europe.Range.–Frasnian UD I-J to Famennian UD II-C. This

group is first known in the German Büdesheimer Schiefer

from levels probably corresponding to the former upper cor-datum Zone (I gamma; now UD I-J), the source of the typespecies, Crassotornoceras ausavense crassum. The new materialfrom the Point Breeze Goniatite Bed of the Angola Shale isprobably of the same age. Crassotornoceras belgicum, fromthe Assise de Matagne of Belgium, is from the holzapfeliZone equivalents, perhaps UD I-K, hence later in age.There are also records of the genus in the lower Famenn-

ian. Grüneberg (1925: 67, pl. 1, figs 11, 11a, 11b) referreda specimen from the Cheiloceras Stufe of the Herzkamp Syn-cline to Steininger’s species Goniatites ausavensis Steiniger(1853: 43, pl. 1, figs 9, 9a) but his figure seems closer to thespecimen figured by Frech (1902a: pl. 4, fig. 9b) from theCheiloceras Stufe at Nehden, Germany. Becker (1993a) hasrevised the lower Famennian occurences.

Crassotornoceras aff. crassum (Matern, 1931a)Pl. 33, Figs 9-20; Text-figs 53E-F; Table 27

aff. Tornoceras ausavense crassumMatern, 1931a: 27, pl. 3, figs 14a-b.

Type material.–The holotype of Crassotornoceras crassumwas designated by Matern as the figured specimen (Senck.Mus. XI 342a) from Büdesheim (UD I-J). The holotype is7.6 mm in diameter with a whorl height in excess of widthand showing four constrictions. Because there is no evidenceof a small saddle centered on the seam, reference to Loboto-rnoceras ausavense is inappropriate.

New York material.–Seven specimens from the AngolaShale (NYSM 12010-12015, 3229/17), none exceeding11.6 mm in diameter and all preserved as partial calciticmolds in gray mudrock.

Dimensions.–See Table 27.Description.–Inner whorls at 3-5 mm diameter involute,

globular, with well-rounded umbilical wall and subcircularwhorl section. Whorl section becoming more compressedwith well-rounded umbilical wall and convexly flat flanksthat converge to well-rounded venter. Ornament withmarked constrictions between 3 and 11 mm diameter, num-bering four per whorl in early stages but reaching seven oreight in outer stages of body chamber. Constrictions follow-

Table 27. Biometric data for Crassotornoceras aff. crassum (Matern, 1931) from the Angola Shale.


NYSM 12010 4.0 max. — — — — —3.2 2.3 — ca. 0.4 — 0.13

NYSM 12012 3.5 max. — — — — —3.2 2.3 — 0.3 — 0.09

NYSM 12015 6.6 max — — — — —5.3 3.8 — ca. 0.0 — —



ing course of growth lines (Text-figs 53E-F) and bettershown on internal molds than on outside of recrystallizedtests where course marked only by shallow concavity, if at all.On internal molds, constrictions scarcely developed on um-bilical wall or shoulder, deep across flanks, rather shallowjust dorsal of ventrolateral salient and again deeper in ventralsinus. Growth lines showing salient on lowest flanks, and inearly whorls a shallow sinus on mid-flanks, with lines pro-jecting slightly to weak ventrolateral sinus and shallow ven-tral sinus; in later whorls, lateral sinus relatively deeper andventrolateral sinus significantly more projecting, almost sub-angular.Sutures, seen only on smaller specimens (NYSM 12012),

with umbilical lobe; generally convexly and backwardly di-rected ventral suture with weak asymmetrical lateral saddle.Ventral lobe small, passing convexly to rounded ventrolateralsaddle.

Discussion.–This form is very close to Crassotornocerascrassum, but the constrictions in that species continue overthe umbilical shoulder and, even at a larger size, are less fre-quent. One of the paratypes of Aulatornoceras loeschmanniFrech [1902a: 49, pl. 4 (5), fig. 9a; Berlin Mus. C 475/1] ismore similar, but this specimen shows constrictions passingover the umbilical shoulder also and, as with C. crassum,more lingulate sutures of a type not apparent in the smallerspecimens from New York that show sutures.

Distribution.–Point Breeze Goniatite Bed (or approxi-mate equivalent), Angola Shale (Erie and Wyoming coun-ties). NYSM 12010 and 12011, Hampton Brook (ErieCounty; Loc. 74/6a); NYSM 12012-12013 and 3229/17,Cazenovia Creek (Erie County; Loc. 75/6); NYSM 12014,Johnson Creek (Wyoming County; Loc. 81/6); NYSM12015, Relyea Creek (Wyoming County; Loc. 82/6).

Range.–Frasnian: Neomanticoceras Genozone UD I-J.Regional Zone of Sphaeromanticoceras rhynchostomum (22b).Conodont Zone MN 11-?12.

Genus EPITORNOCERAS Frech, 1902aType species.–Goniatites mithracoides Frech, 1887.Diagnosis.–Tornoceratids with oxyconic form in adult

or, in some, earlier stages; with or without acute ventrolateralsaddle. Otherwise like Tornoceras.

Included species.–There are three quite different oxyconicstocks derived from “normal” Tornoceras of the T. uniangu-lare/simplex stock. The groups are:

Group 1. Gen. nov. Small oxycones with Tornoceras uni-angulare-style sutures, but with open umbilicus and“pinched” and acute ventral margin. Seen only from a spec-imen in the collection of G. Kloc (University of Rochester)from the Wanakah Shale (Hamilton Group), northwest ofBethany Center (Genesee County).

Group 2. Epitornoceras. This has always been interpretedas an oxyconic genus, but reillustration of the type species,Epitornoceras mithracoides, by House (1978: pl. 10, figs 2-3;Berl. Mus. c469) has shown that species not to be truly oxy-conic, only laterally compressed with a tendency to becomesubacute in larger whorls (shown by the second, unillus-trated specimen in the Museum für Naturkunde, Berlin,Berl. Mus. C470). Nor is the ventrolateral saddle as sharplyacute as shown in many illustrations. Nevertheless it seemsprobable that E. mithracoides belongs to the early part of thestock that gives the true oxycones with sharp ventrolateralsaddles, such as E. peracutum, which occur in the late Givet-ian Pharciceras Stufe, and probably only in the lower parts.Included are: E. mithracoides (Frech, 1887: 30, pl. 2, fig. 1;House, 1978: pl. 10, figs 2-3); E. peracutum (Hall, 1879:pl. 69, fig. 8, pl. 74, fig. 13: holotype, NYSM 4091, figuredby House, 1965: text-fig. 13C); E. aff. peracutum (figured byHouse, 1978: 60, pl. 6, figs 7-8, 11-12, text-fig. 7B); E. aff.peracutum (figured by Harris, 1899: pl. 6, fig. 35; refiguredby House, 1965: text-fig. 13H, pl. 8, fig. 74, NYSM/CU39652).

Group 3. Oxycones are again derived from the conser-vative Tornoceras stock in the early Nehdenian. These havebeen named Oxytornoceras (Becker, 1993a) with a typespecies, by original designation, of T. acutum (Frech, 1902a:47, pl. 3(2), fig. 17). A sequence of forms giving a clear linkfrom typical Tornoceras to oxyconic forms with an acute ven-ter has been elucidated by the late Dr. H. Makowski in col-lections from the early Famennian of the Holy CrossMountains of Poland. These might be quite homeomorphicwith the true Epitornoceras in that the ventrolateral saddlecan, unusually, become acute.Material of Group 2 is only available for description

from New York. The range and distribution data givenbelow apply only to Group 2.

Discussion.–New York specimens agree in the oxyconicform with the usual interpretation of the genus, but itshould be emphasized that the comparison with the typespecies is based on the similar large, lingulate symmetricallateral lobes and not on whorl form. The earliest of theknown New York representatives (NYSM/CU 42401) stillshows a marked asymmetry in the lateral lobe (Text-fig.57F). The holotype of E. peracutum (Text-fig. 57A; NYSM4091) and the other New York andWest Virginia specimensare all closely united in sutural form, but there is variationin depth of lateral lobe. New York is the only place in theworld where there is evidence bearing on the time-successionof the group. NYSM/CU 40105, from the Geneseo Shale atHubbard Quarry (Seneca County; Loc. 27b), is the largestknown tornoceratid from North America, if not the world.The original figures of Oxytornoceras acutum show the

growth lines well and they differ from the large New Yorkspecimen described here (Text-fig. 57E) in not having awell-developed salient on the lower flanks. This could beone means of distinguishing the early Famennian groupfrom the true Epitornoceras.

Distribution.–North America (New York and West Vir-ginia) and Europe (Germany).

Range.–Givetian MD II to Frasnian UD I-A. The earlyFrasnian oxyconic tornoceratids correctly referred to Epi-tornoceras occur in Germany only in the Pharciceras Stufe(Frech, 1902a, b). In New York, forms assigned to the genusappear in the upper part of the Geneseo Shale (GeneseeGroup) and records continue to at least the middle PennYan Shale, thus they range above the Givetian-Frasnian

boundary. The specimen from the Harrell Shale ofWest Vir-ginia (House, 1978) with ?Pharciceras galeatumWedekind,1917, probably correlates to the earlier part of the New Yorkrange.

Epitornoceras cf. mithracoides (Frech, 1887)Text-fig. 57F

cf. Goniatites mithracoides Frech, 1887: 30, pl. 2, fig. 1.cf. Tornoceras (Epitornoceras) mithracoides. Frech, 1902a: 51;

1902b: 172 (text-fig.); 1913: 19.Epitornoceras peracutum (pars). House, 1965: 118-119.Epitornoceras mithracoides. House, 1978: 59, pl. 10, figs 2-3.

Type material.–Frech’s original material is in the Museumfür Naturkunde (Berl. Mus. c469, 470) and one specimenhas been refigured by House (1978).

New York material.–One large crushed specimen fromthe Geneseo Shale (NYSM/CU 42401) preserved in darkgray shale.

Dimensions.–NYSM/CU 42401: D = ca. 200 mm;WH= 120 mm; Wh = 64 mm; UW = 0 mm.

Description.–The body chamber seen to 340°; shellcrushed; shell form not discernable. Involute with closedumbilicus. Suture as shown in Text-fig. 57F. Aperture notseen but evidence suggesting salient low on flanks; anotherinferred in usual ventrolateral position.

Discussion.–This specimen is compared to Epitornocerasmithracoides because of the large size and the similar, slightlyasymmetric large lingulate lateral lobe that distinguishes itfrom the later form, E. peracutum. Epitornoceras mithracoidesseems to have its closest analog among the mainstreamtornoceratids, with Tornoceras arcuatum, the type materialof which is from the much later Squaw Bay Limestone ofMichigan. Rather similar forms, recorded as T. cf. arcuatum,are already known in the Tully Limestone (House, 1965,and herein), but these have narrower lateral lobes and morehighly arched and symmetrical latero-umbilical saddles thanE. mithracoides. Presumably something like T. cf. arcuatumgave rise to Epitornoceras as indicated previously (House,1965: text-fig. 1, a figure that was not intended to implythat the Michigan T. arcuatum belonged to the Tully level).

Distribution.–Upper Geneseo Shale (Genesee Group),Lodi Glen (Seneca County; Loc. 27).NYSM/CU 42401 was collected by J. W. Wells in Lodi

Glen (Loc. 27; his loc. 111g), 12.2 m (40 ft) below the topof the Geneseo Shale and approximately 24 to 27.5 m (80-90 ft) above the Tully Formation.

Range.–Givetian MD III (?C-?D). Regional Zone of Epi-tornoceras peracutum (15a), but occurring before the zonalspecies.

Text-fig. 57. Sutures, cross section, and outline of Epitornocerasspp. from the Genesee Group in New York, the Harrell Shale inWest Virginia, and Oberscheld, Germany. A. E. peracutum (Hall,1879), NYSM 4091, suture of the holotype at 41 mm whorlheight, Ithaca Shale and Sandstone, Ithaca, Tompkins County, X0.5. B-C. E. aff. peracutum. B. USNM 186052, reversed suture ofa specimen figured by House (1978: 34) from the Harrell Shale,near Landes Post Office, West Virginia (see Weary & Harris,1994), X 0.5. C. NYSM/CU 39652, suture of a specimen figuredby Harris (1899: pl. 6, fig. 35) at ca. 80 mm diameter from theIthaca Shale and Sandstone, Ithaca, Tompkins County, X 0.5. D-E. E. cf. peracutum. D. NYSM/CU 369, cross section based on asolid specimen from the upper Geneseo Shale or lower Penn YanShale, southern side of Fir Tree Point (Loc. 25), Seneca Lake,southeast of Rock Stream, western side of Seneca Lake, YatesCounty, X 0.63. E. NYSM/CU 40105, outline (drawn by J. W.Wells) showing the apertural margin of a specimen reaching ca.240 mm diameter from the upper Geneseo Shale at HubbardQuarry (Loc. 27b), Seneca County, X 0.13. F. E. cf. mithracoides(Frech, 1887), NYSM/CU 42401, suture of a specimen from 12.2m (40 ft) below the top of the Geneseo Shale in Lodi Glen (Loc.27), Seneca County, X 0.5. G. E. mithracoides, BM c469, reversedsuture of one of the cotypes of Frech (1887a) from Grube Eibach,Oberscheld, Germany, X 0.5.


Epitornoceras peracutum (Hall, 1876) GroupPl. 34, Fig. 8; Text-fig. 57A; Table 28

Goniatites peracutusHall, 1876: pl. 69, fig. 8, pl. 74, fig. 13; 1879:463, pl. 69, fig. 8, pl. 74, fig. 13.

Tornoceras peracutum. Clarke, 1898, 1899a, b: 118, text-fig. 96.Tornoceras (Epitornoceras) peracutum. Frech, 1913: 19.Tornoceras (Tornoceras) peracutum.Miller, 1938: 155-156, pl. 31,

figs 1-2.Epitornoceras peracutum. House, 1965: 118-119, text-fig. 13c, pl.

8, figs 73-74.

Type material.–The holotype (NYSM 4091) has been de-scribed by Miller (1938) and House (1965).

New material.–Other specimens from the Geneseo Shaleconforming to the species include: NYSM/CU 40105 (Text-fig. 57E), an extremely large, crushed specimen, andNYSM/CU 369, a solid specimen (Text-fig. 57D, Pl. 34,Figs. 6-7), both referred to Epitornoceras cf. peracutum, andless well preserved material and specimens seen in the fieldbut uncollectible. NYSM/CU 39652, the specimen figuredby Harris (1899: pl. 6, fig. 35), is refigured here (Text-fig.57C, Pl. 34, Fig. 5) and is referred to E. aff. peracutum.

Dimensions.–See Table 28.Discussion.–In view of the illustrations given here, much

additional comment is unnecessary. The cross section ofNYSM/CU 396 (Text-fig. 57D) shows the progressive onsetof oxyconic form that is achieved after approximately 25mm diameter. There is some variation in the depth of thelateral lobe, and NYSM/CU 39652 (Text-fig. 57C) is distin-guished from the holotype by the lesser depth of the laterallobe.

Distribution.–Upper Geneseo Shale to middle Penn YanShale and equivalents in the Ithaca Formation (GeneseeGroup; Tompkins County), westward to Livingston County.Harrell Shale (Grant County), West Virginia.NYSM/CU 40105, the very large specimen from at or

near the top of the Geneseo Shale at Hubbard Quarry(Seneca County; Loc. 27b).NYSM/CU 369, from upper Geneseo Shale or lower

Penn Yan Shale reported from “between Rock Stream andRock Landing,” southern side of Fir Tree Point, Seneca Lake(Yates County; Loc. 25).The source of the holotype (NYSM 4091) and

NYSM/CU 39652 is only given as Ithaca in TompkinsCounty, and it can only be presumed that they are fromlower levels of the Ithaca Formation. The rare specimens ofEpitornoceras known elsewhere from above the GeneseoShale are from well up in the Penn Yan Shale [3846/1,Taunton Gully (Loc. 17/4); 3853/4, Fall Brook-Dewey Hill(Livingston County; Loc. 19/5)]. These Penn Yan Shalespecimens from Livingston County are poorly preserved.USNM 186052, Epitornoceras aff. peracutum, figured by

House (1978: 60, text-fig. 7B, pl. 6, figs 7-8, 11-12; suturerefigured here, Text-fig. 57B), is from the Harrell Shale, 0.8km (0.5 mi) south of Landes Post Office (Grant County),West Virginia.

Range.–Givetian MD III (?C-?D) to Frasnian UD I-A.Regional Zone of Epitornoceras peracutum (15a) to Ponticerasperlatum (15b).

Genus LOBOTORNOCERAS Schindewolf, 1936Type species.–Goniatites ausavense Steininger, 1853, by

original designation (Schindewolf, 1936: 689), a form fromBüdesheim, Germany; topotype figured by Schindewolf(1936) and neotype of House (1978), SM H9932, hereinText-figs 58C, F.

Diagnosis.–Tornoceratids with closed or nearly closedumbilicus, well-rounded compressed to subplatyconic form,rarely with ventrolateral furrows; some with prominent lat-eral constrictions. Suture as in Tornoceras but with additionalsaddle centered on umbilical seam (EAL’UD). Growth linesbiconvex.

Included species.–A neotype for Lobotornoceras ausavense(Steininger, 1853: 40, pl. 1, fig. 11) was selected and figuredby House (1978: pl. 9, figs 8, 14; Sedgw. Mus. H 9932).Additionally there is L. hassoni House (1978: 59, pl. 8, figs6-7, 10-11, text-figs 11E-G; holotype USNM 239891,paratypes 239892-239896, 240505) and L. aff. hassoni de-scribed herein. Famennian species formerly grouped herehave been assigned to the genera Falcitornoceras and Exo-tornoceras by Becker (1993a).

Discussion.–This genus appears to arise from Tornocerass. s. by the addition of a small saddle centered on the um-bilical seam. This happened at least twice but the Frasnianspecies show a distinctive, broad, subsymmetrical laterallobe, whereas later Famennian homeomorphs show more

Table 28. Biometric data for Epitornoceras peracutum (Hall, 1876) from the Geneseo Shale and Ithaca Shale and Sandstone.


NYSM 4091 — — 41- — 0 — —(holotype)

CU 40105 ca. 240 — ca.142- ca. 74 0 — —NYSM 369 75 18 45- — 0 0.40 —


sharply folded sutural elements.Distribution.–North America and Europe.Range.–Frasnian: UD I-C to UD I-I/J. In North Amer-

ica: Frasnian: UD I-C. Lobotornoceras hassoni is recordedfrom a styliolinid limestone, possibly close to the level of theGenundewa Limestone, in the Harrell Shale in West Vir-ginia (House, 1978). Lobotornoceras aff. hassoni is describedherein from the West River Shale of New York. One speci-men of the homeomorphic Falcitornoceras aff. bilobatum(Wedekind, 1908) is known from the Famennian in theNorthwest Territories of Canada (House & Pedder, 1963).

Lobotornoceras aff. hassoni House, 1978Pl. 34, Figs 1-4; Text-figs 58A-B; Table 29

aff. Lobotornoceras hassoniHouse, 1978: 59, pl. 8, figs 6-7, 10-11;text-figs 11E-G.

Type material.–The holotype of Lobotornoceras hassoni(USNM 239891; Text-fig. 58E) and paratypes (USNM239892-239896, 240505) are from a styliolinid-rich nodulein the Harrell Shale, from 4.1 mi (6.6 km) south of LandesPost Office (Grant County), West Virginia.

New York material.–Two specimens only from the WestRiver Shale in Ontario County: NYSM 12029, a crushedphragmocone preserved as an incomplete pyritic internalmold, from Seneca Point (Loc. 23a/3); NYSM 12030, asolid, incomplete, pyritized internal mold, fromWhetstoneBrook (Loc. 26a/1).

Dimensions.–See Table 29.Description.–The larger specimen (NYSM 12029) shows

approximately 18 sutures in the last half whorl. Shell formdifficult to determine due to crushing but the maximumwhorl width appears to lie approximately one third of thedistance from the umbilicus to the venter; the flanks slopeevenly to the umbilicus without any shoulder; the venter ap-pears to be abruptly rounded and could have a flattening.Suture as illustrated in Text-fig. 58A. The smaller specimen(NYSM 12030; Text-fig. 58B) indicates a shell form againwith the maximum whorl width approximately one third ofthe distance from the umbilicus to the venter with no um-bilical shoulder; toward the venter the flanks converge con-vexly and there is a well-rounded ventrolateral shoulder anda flatly rounded venter; suture as illustrated inText-fig. 58B.

Discussion.–Sutural form of the larger of these specimensapproaches the holotype of Lobotornoceras hassoni but bothspecimens differ in the absence of evidence of a well-rounded form and flattish venter. Both specimens differfrom L. ausavense in showing no constrictions, and from Fa-mennian homeomorphs in the form of the suture.

Distribution.–Middle to upper West River Shale (Gene-see Group; Ontario County).

Range.–Frasnian: TimanitesGenozone UD I-C. RegionalZone of Koenenites beckeri (17b). Conodont Zone MN 4.

Tribe FALCITORNOCERATINI Becker, 1993aDiagnosis.–Tornoceratidae similar to the Tornoceratini

Table 29. Biometric data for Lobotornoceras aff. hassoni House, 1978, from the West River Shale.


NYSM 12029 7.4 — 4.8 ? 0.3 — 0.04NYSM 12030 6.7 max. — — — — —NYSM 12030 — 2.1 3.24 ? 0.3 0.65 —

Text-fig. 58. Sutures and cross section of Lobotornoceras spp. fromNew York,West Virginia, Northwest Territories, and Germany. A-B. L. aff. hassoni House, 1978. A. NYSM 12029, reversed suturebased on a specimen from the upper West River Shale on SenecaPoint Creek (Loc. 23a/3), X 4.6. B. NYSM 12030, suture andwhorl section of a specimen from the upper West River Shale onWhetstone Brook (Loc. 26a/1), near Honeoye, Ontario County;suture at 3.2 mm whorl height and cross section at 3.4 mm whorlheight, X 4.6. C. L. ausavense (Steininger, 1853), suture of a spec-imen from Büdesheim, Germany, figured by Schindewolf (1936),reversed for comparison; magnification not stated. D. Falci-tornoceras aff. bilobatum (Wedekind, 1908), GSC 16955, suture at34 mm diameter figured by House & Pedder (1963: 529) fromnear Carlson's Lake, Northwest Territories, X 0.8. E. L. hassoni,USNM 239891, suture of the holotype at 5.8 mm whorl heightfrom the Harrell Shale south of Landes Post Office, West Virginia(see Weary & Harris, 1994, for locality details), X 4. F. L.ausavense, SM H 9932, suture of the neotype from the Büde-sheimer Schiefer at 210 m south-southeast of Büdesheim Church,Germany, X 4.6.


but commonly with ventrolateral furrows in earliest stagesand falcate early ornament, and with lateral constrictions.Some with additional umbilical saddle.

Discussion.–In this group, Becker (1993a, 1995) in-cluded Phoenixites, Falcitornoceras House & Price, 1985[type speciesGoniatites (Tornoceras) subundulatus var. falcataFrech, 1887], and Gundolficeras Becker, 1995 (type speciesLobotornoceras bicaniculatum Petter, 1959), the last twocommonly developing a saddle on the umbilical seam.Here we follow the taxonomy of Becker (1993a, 1995),

but with some misgiving because detailed ontogenies havenot been described for many of the included forms. Mostof the many species of Tornoceras have early stages that arequite unknown and their assignment is put in questionwhen a genus (Phoenixites) is erected based only on charac-ters of the early stages.

Distribution.–Worldwide. In New York, represented byPhoenixites concentricus in the Gowanda Shale (CanadawayGroup).

Range.–Probable worldwide range: Givetian MD II toFamennian UD VI. In New York: Famennian: UD II-C.

Genus PHOENIXITES Becker, 1993Type species.–Tornoceras frechiWedekind, 1918, by orig-

inal designation (Becker, 1993a: 198).Diagnosis.–Like Tornoceras but with ventrolateral grooves

in the early stages and some with constrictions.Included species.–See Becker (1993a, 1995).Distribution.–North America, Europe, Russia, and

North Africa.Range.–Upper Givetian MD III-A to Famennian UD II-

C.

Phoenixites concentricus (House, 1965)Pl. 29, Figs 1-15; Text-figs 54M-O, 59A; Table 30

Tornoceras (Tornoceras) concentricumHouse, 1965: 113-115, text-

fig. 14, pl. 9, figs 84, 86-88, pl. 10, figs 101-110, pl. 11, fig.138.

Phoenixites concentricus. Becker, 1993a: 198.

Type material.–The holotype [NYSM 11963 (6679D)]and nine other specimens from the Gowanda Shale were de-scribed by House (1965).

New material.–Eight additional specimens are available(listed below).

Dimensions:–See Table 30 and Text-fig. 59A.Discussion.–The new material adds extra statistical data

that are plotted in Text-fig. 59A for comparison with otherknown tornoceratids. Other details remain as given in theprimary description in which details of the earliest stageswere given. The diagnostic suture of the species, with its ex-ceedingly steep ventrad face to the latero-umbilical saddleand the sharp turn over the ventral crest of that saddle inadult stages, is illustrated in Text-figs 54M-N. There is nonew evidence regarding sexual dimorphism speculated uponearlier (House, 1965: 115). Too little is known of the earlystages of most Tornoceratidae for this species to be separatedfrom the Tornoceratini with any confidence.

Distribution.–Corell’s Point Goniatite Bed, GowandaShale (Canadaway Group; Chatauqua, Cattaraugus, andWyoming counties).Only known from the Corell’s Point Goniatite Bed of

the Gowanda Shale. Material listed by House (1968) andNYSM/CU 3292/1, 6, 14, Corell’s Point on Lake ErieShore (Chatauqua County; Loc. 97); NYSM 12052, Big In-dian Creek (Cattaraugus County; Loc. 101); NYSM 12053and 12054, and NYSM/CU 3278/20, Cattaraugus Creek(Cattaraugus County; Loc. 104); NYSM 16565 and 16566(Pl. 29, Figs 11-12), Java (Wyoming County; Loc. 109).Poorer material of this species has been noted at almost everyoutcrop of the Corell’s Point Goniatite Bed inland of LakeErie.

Range.–Famennian: Cheiloceras (Cheiloceras) Genozone

Table 30. Biometric data for Phoenixites concentricus (House, 1965) from the Gowanda Shale. See Text-fig. 59A for graph.


NYSM 11963 20.5 9.4 12.7-7.5 ca. 1.0 0.74 0.05(holotype)

NYSM 12053 ca. 39.0 11.8 22.6-12.5 ca. 0 0.52 —NYSM 12054 ca. 33.9 ca. 13.5 21.3-12.5 ca. 0 0.63 —NYSM 12052 28.8 12.2 18.1-9.7 ca. 0 0.67 —

19.2 9.3 ca. 11.2- — ca. 0 0. 83 —NYSM 11965 18.0 7.9 10.5- — ca. 0 0.75 —NYSM 12053 16.3 7.8 ca. 9.8- — ca. 0 0.79 —NYSM 11964 13.3 6.7 8.0- — ca. 0 0.84 —USNM 137668 12.3 5.0 — 0 — —NYSM 11965 1.5 1.01 0.7- — 0.23 1.44 0.15


Text-fig.59.Graphsshowingontogenyof

Phoenixites,Truyolsoceras,andCheiloceras,allbasedonmaterialfromtheCorell'sPointGoniatiteBed,GowandaShale,inwesternNew

York.SeeTables30and34-35forbiometricdata.A.P.concentricus(House,1965),asTornoceras

concentricum

,basedonmateriallistedhereinandbyHouse(1965)from

Corell's

Point(Loc.97),BigIndianCreek(Loc.101),andCattaraugusCreek(Loc.104),ChautauquaandCattarauguscounties.B.T

.bicostatum(Hall,1843),asAulatornocerasbiocosta

tum,

basedonmateriallistedhereinandbyHouse(1962)from

Corell'sPoint(Loc.97)andWalnutCreek(Loc.99b),ChautauquaCounty.C.C

.(Cheiloceras)am

blylobum

(G.&

F.Sandberger,1851),basedonmateriallistedhereinandbyHouse(1962)from

Corell'sPoint(Loc.97),ChautauquaCounty.


UD II-C. Regional Zone of C. amblylobum (25).

Tribe AULATORMOCERATINI Becker, 1993aGenus AULATORNOCERAS Schindewolf, 1922Type species.–Goniatites aurisQuenstedt, 1846, by origi-

nal designation (Schindewolf, 1922: 188). Neotype chosenby House & Price (1985: pl. 17, figs 1-4; SMH9942) fromBüdesheim.

Diagnosis.–Tornoceratidae with ventrolateral furrows,commonly flat-sided with tabular venters and closed ornearly closed umbilicus in early stages to subinvolute inadult. Ornament forming strong ribs, or with bunching ofprominent growth lines or radial bands; in some withgrowth-line festoons on venter increasing in frequency inadult; most with periodic constrictions.

Discussion.–Miller (1938: 142) drew attention to the twogroups of morphologies then included within Aula-tornoceras. He distinguished between a group with open anda group with closed umbilici. There is clearly a wide rangeof very different forms included within the genus at present,

but until European species are more precisely defined, bothtaxonomically and stratigraphically, little progress will bemade in the elucidation of the evolution of the group. Itseems possible to recognize three groups of aulatornoceratidsas follows:

Group A. These include the type species and are charac-terized by a closed or nearly closed umbilicus and ornamentwithout very strong ribs or constrictions but with adults thatare rather smooth-sided although the growth lines can beprominent on the flanks and commonly form festoons onthe venter. Included species: Aulatornoceras auris (Quenst-edt, 1846: 64, pl. 3, figs 7a-c); ?A. auris bickensis (Wedekind,1918: 137; Matern, 1931a: 31); A. eifliense (Steininger,1849: 27; 1853: 43, pl. 1, figs 3, 3a, but not 2, 2a); A. ei-fliense posterior Becker (1993a: 216, text-figs 77e-f, 78d, pl.9, figs 7-10); A. sandbergeri (Foord & Crick, 1897: 112 proG. & F. Sandberger, 1851: pl. 10, figs 17, 17a; notGoniatitesundulatus Brown, 1841); and A. lepiferum Becker (1993a:215, text-figs 77c, 78b, pl. 8, figs 11-13, pl. 9, fig. 1).

Group B. These are forms with markedly open umbilici

Text-fig. 60. Sutures and cross sections of Truyolsoceras and Aulatornoceras spp. from the Gowanda Shale and Angola and Hanover Shales inwestern New York. A-C. T. bicostatum (Hall, 1843), from the Corell's Point Goniatite Bed, Gowanda Shale, Corell's Point (Loc. 97), Chau-tauqua County. A. Suture at ca. 20 mm diameter based on USNM 137662, X 2. B. NYSM 11958 (D.1363), suture at 12.5 mm diameter,X 3.2. C. NYSM 11957 (D1364), suture at 4.5 mm diameter, X 3.2. D. A. paucistriatum (d'Archiac & de Verneuil, 1842), NYSM 12094,suture at 9 mm diameter of a specimen from the Point Breeze Goniatite Bed in in the Angola Shale, Cazenovia Creek (Loc. 75/6), ErieCounty, X 4. E. A. rhysum (Clarke, 1898), NYSM 4092, suture and growth line of the lectotype at 5 mm diameter, based on a specimensaid to come from the Hanover Shale at Java, Wyoming County, X 9.2. F. A. paucistriatum, NYSM 12094, cross section of same specimenas in D, X 4. G. A. auris (Quenstedt, 1846) Group, cross section based on NYSM 12113 from the lower Angola Shale in Hampton Brook(Loc. 74/3), Erie County, X 4.4. H. T. bicostatum, NYSM 11959 (= NYSM 6679), same location as A-C, cross section at 18.7 mm diameter,X 3.2.



in early whorls, and open umbilici in later whorls, and thatare extremely heavily ribbed in the early whorls althoughribbing declines in prominence on the outer whorls. In-cluded species: Aulatornoceras paucistriatum (d’Archiac & deVerneuil, 1842: 339, pl. 25, figs 8, 8a, 8b); A. rhysum(Clarke, 1898: 121, text-fig. 100, pl. 8, fig. 14; NYSM4092); and A. serriense Becker (1993a: 213, text-figs 77a-b,78a, pl. 8, figs 14-18).

Group C. These are closed umbilicate, rather smooth-surfaced aulatornoceratids with periodic constrictions. In-cluded species: Aulatornoceras constrictum (Steininger, 1853:43, pl. 1, fig. 9); A. auriformeOppenheimer (1916: 174, pl.1, fig. 2; see Becker, 1993a); A. keyserlingiMüller (1956: 49pro Keyserling, 1846: 277, pl. 12, fig. 3; not Goniatites cinc-tumMünster, 1843); and A. loeschmanni (Frech, 1902a: 49,pl. 4 (5), figs 9a-c)

Discussion.–The first two groups are well represented inthe New York State Devonian and the third group not at all.Some use is made of European names in the account thatfollows because there can be little doubt as to the close rela-tionship of the forms concerned, but this is bedeviled by thefact that, despite wide use of the names in Germany, few ofthe German type specimens have been figured photograph-ically since they were originally described.

Distribution.–North America, Europe, Russia, NorthAfrica, and Western Australia.

Range.–Frasnian: UD I-I to Famennian UD II-D.

Aulatornoceras auris (Quenstedt, 1846) GroupPl. 31, Figs 2-3; Text-fig. 60G

Goniatites auris Quenstedt, 1846: 64, pl. 3, figs 7a-c.

Type material:–The type material of this species has notbeen redescribed or figured since its first description. Thespecimen figured by Quenstedt was said to come fromBüdesheim, Germany, and a large number of horizons thereyield aulatornoceratids. Despite the fact that the species isthe type-species of the genus, it is not critically determinableby modern standards. It is used here in a wide sense.

New York material.–Generally poorly preserved largespecimens are included here (NYSM 12113-12118) fromthe Angola and Hanover Shales, none of which are com-plete. Most show evidence of the body chamber only, all arepreserved in mudrock with some shell replacement by cal-cite, and some have crystalline calcite infilling.

Description.–This material does not warrant detailed de-scription, but illustrations are given (Pl. 31, Figs 2-3, Text-fig. 60G). One solid specimen (NYSM 12113) shows theproportions of the inner and outer whorls (Text-fig. 60G);other specimens are less well preserved. Most specimens

show the development of ventral festoons (Pl. 31, Fig. 3).None of the specimens shows the suture.

Discussion.–This is a “waste-basket” group used for largeaulatornoceratids with nearly closed umbilici (when seen)and united by the development of festoon-like grooves pe-riodically on the venter that follow the growth line course.This is a feature of large specimens generally referred toAulatornoceras auris from Büdesheim. The possibility thatouter whorls of A. paucistriatum, A. rhysum, and A. eifleinseare included here cannot be discounted, although the lastwould be unlikely. The one cross section (Text-fig. 60G)suggests a different early stage than these species. From theGowanda Shale, Truyolsoceras bicostatum does not appear todevelop festoons and T. clarkei is more widely umbilicate.The alternative to using the open nomenclature followed

here would be to erect new names, but this would be inap-propriate at present. Taxonomic revision and detailed de-scription of the European species, particularly fromGermany, are needed.

Distribution.–Angola Shale and Hanover Shale (Erie andWyoming counties).Type material from Büdesheim, Gemany. This species

name has been used widely in Germany and internationallyfor Frasnian aulatornoceratids. The exact source of the ma-terial of Quenstedt from Büdesheim is not known, but isprobably from the main locality southeast of the church,and hence a level in the late cordatum Zone is most probable,but the species could well range higher.Becker et al. (2000: 78, fig. 4) reported Aulatornoceras

cf. auris from the Lyaiol Formation in Timan, Russia (UDI-I).In New York, the records are from the Angola Shale

(below the Point Breeze Goniatite Bed) and lower HanoverShale, but not also from upper Rhinestreet Shale as stated byHouse & Kirchgasser (1993: 276, 279, fig. 7) and Becker &House (2000: 129). Angola Shale: NYSM 12113, HamptonBrook (Erie County; Loc. 74/3); NYSM 12114 and 12115,Sheldon Creek (Erie and Wyoming counties; Loc. 78/4b).Hanover Shale: NYSM 12117 and 12118, Beaver MeadowCreek, Java (Wyoming County; Loc. 92/3).

Range.–In New York: Frasnian: Neomanticoceras Geno-zone (UD I-J) to Archoceras Genozone (UD I-K). RegionalZone of Sphaeromanticoceras rhynchostomum (22b), Delph-iceras cataphractum (23). Conodont Zone MN 11-12.

Aulatornoceras eifliense (Steininger, 1849)Pl. 31, Figs 7-13, 16-20; Table 31

Goniatites eifliensis Steininger, 1849: 27; 1853: 43, pl. 1, figs 3,3a.

Goniatites retrorsus var. undulatus G. & F. Sandberger, 1851: pl.10, fig. 9.

Tornoceras eifliense.Wedekind, 1918: 137.Aulatornoceras eifliense. House & Kirchgasser, 1993: 276.Aulatornoceras eifliense eifliense. Becker, 1993a: pl. 9, figs 4-6.

Type material.–Steininger’s material has neither been re-described nor rediscovered. Wedekind (1917) selected oneof the specimens figured by Steininger (1853: pl. 1, fig. 3)and this could be construed as selection of a lectotype. Butthis has been overtaken by the selection as a neotype byBecker (1993a: 370) of a specimen in the Wiesbaden Mu-seum figured by the Sandberger brothers (1851: pl. 10, fig.19) from Büdesheim.

NewYork material.–Seven specimens from the CashaquaShale (NYSM 12105-12111) including some earliest stagespreserved in pyrite but larger specimens preserved as bariticreplacements of the shell.

Dimensions.–See Table 31.Description.–Inner whorls with closed umbilicus from

approximately second whorl. Whorl form at first showingsignificant lateral compression, with convex flanks, ventro-lateral furrows, and convex ventral band conforming to totalwhorl outline (NYSM 12105; Pl. 31, Figs 10-11). Whorlwidth increasing relatively and by largest diameter seen(NYSM 12107; Pl. 31, Figs 18-20), with whorl form dif-fering from Büdesheim types in being less compressed andmore inflated. Sutures showing V-shaped ventral lobe andasymmetric lateral lobe in early whorls at approximately 5mm diameter (NYSM 12105; Pl. 31, Fig. 11); suture notseen at larger diameters. Shell surface generally smooth,without constrictions, ribs, or festoons on venter. Ornamentconsisting of growth lines alone, with striae forming lateralsinus, passing forward to form lappet centered in ventrolat-eral groove, and passing back to U-shaped ventral sinus.

Discussion.–Wedekind used the species name Tornoceraseifliense for specimens like Aulatornoceras auris but with aclosed umbilicus and without constrictions. That is the sensein which it is used here and conforms to the neotype selected

by Becker (1993a) except that it is further restricted to spec-imens showing the same evidence of a laterally compressedform as indicated by Steininger’s figure.

Distribution.–Cashaqua Shale (Sonyea Group; Erie,Genesee, and Livingston counties).Specimens include: NYSM 12105 and 12111, Smoke

Creek (Erie County; Loc. 31/5); NYSM 12106, CazenoviaCreek (Erie County; Loc. 32/9); NYSM 12110, MurderCreek (Genesee County; Loc. 34/6); NYSM 12109, BuckRun Creek (Livingston County; Loc. 39/17); NYSM12107, Shurtleff Septarian Horizon, Shurtleff ’s Gully (Liv-ingston County; Loc. 41/1).A poorly preserved specimen (NYSM 12112) from the

Angola Shale, Lake Erie Shore (Farnham Creek; Loc. 71/9a),shows a similarly closed umbilicus and unornamented formbut cannot be placed here with any confidence; it is referredto Aulatornoceras aff. eifliense.

Range.–In New York: Frasnian: Probeloceras Genozone(UD I-E) to ProchoritesGenozone (UD I-F). Regional Zoneof Probeloceras lutheri (19) to Prochorites alveolatus (20).Conodont Zone MN 5-6.

Aulatornoceras aff. eifliense (Steininger, 1849)Pl. 31, Figs 4-6, Pl. 32, Fig. 14; Table 32

aff. Goniatites eifliensis Steininger, 1849: 27; 1853: 43, pl. 1, figs3, 3a.

aff. Aulatornoceras eifliense eifliense. Becker, 1993a: pl. 9, figs 4-6.

NewYork material.–Five specimens from the Rhinestreetand Angola Shales. NYSM 12102-12104 preserved as solidphragmocones in pyrite and mudrock, and NYSM 12116 inmudrock.

Dimensions.–See Table 32.Description.–Small subglobular aulatornoceratids with

closed umbilicus, inflated whorl form, and weak ventrolat-eral furrows. Growth lines (NYSM 12104) with salient cen-

Table 31. Biometric data for Aulatornoceras eifliense (Steininger, 1849) from the Cashaqua Shale.


NYSM 12107 13.5 8.0 — 0 — —NYSM 12106 7.9 ca. 4.7 — 0 — —NYSM 12105 5.1 2.4 — 0 — —

Table 32. Biometric data for Aulatornoceras aff. eifliense (Steininger, 1849) from the Angola Shale.


NYSM 12103 7.1 4.4 — 0 — —NYSM 12104 5.9 3.8 — ca. 0 — —


tered on rounded umbilical shoulder, with broad lateralsinus, lappet in ventrolateral groove, and deep U-shapedventral sinus. Sutures (NYSM 12103) with ventral V-shapedlobe, linguiform lateral lobe, and flat-topped latero-umbil-ical saddle. NYSM 12116 (Pl. 31, Fig. 4) showing distinc-tively paired growth lines on flanks.

Discussion.–These are discriminated from Aulatornoceraseifliense on the basis of the more rotund form; they are alsolater in age.

Distribution.–Upper Rhinestreet and lower Angola Shale(West Falls Group; Erie and Wyoming counties).Rhinestreet Shale: NYSM 12116, Kennedy Gulf

(Wyoming County; Loc. 63/1). Angola Shale: NYSM12103 and 12112, Lake Erie Shore (Farnham Creek) (ErieCounty; Loc. 71/9a); NYSM 12102, Hampton Brook (ErieCounty; Loc. 74/6a); NYSM 12104, Sheldon Creek(Wyoming County; Loc. 78/4b).

Range.–In New York: Frasnian: ?Playfordites Genozone(UD I-I) to Neomanticoceras Genozone (UD I-J). RegionalZone of ?P. cf. tripartitus (?22a) to Sphaeromanticoceras rhyn-chostomum (22b). ?Conodont Zone MN 11 and ?earlier.

Aulatornoceras paucistriatum(d’Archiac & de Verneuil, 1842)

Pl. 32, Figs 2-13; Text-figs 60D, F; Table 33

Goniatites paucistriatum d’Archiac & de Verneuil, 1842: 339, pl.25, figs 8, 8a, 8b.

Type material.-The original material has not been re-described, and it was not located during a search of deVerneuil’s collection in the l’Ecole des Mines in Paris madeby one of us (MRH) in 1958. The original material wasrecorded as from Adorf and Oberscheld, Germany.

New York material.-Several specimens from the AngolaShale (including NYSM 12094-12101 and BMNS E22464), mostly preserved in concretionary mudrock withsome crystalline calcite filling of internal whorls. One spec-imen is a siliceous shell replacement (NYSM 12096).

Diagnosis.–This name is used here for openly umbilicateAulatornoceras conforming to the type illustrations withstrong ribs or grooves on the flanks, especially in the innerwhorls.

Dimensions.–See Table 33.Description.–Small, openly umbilicate aulatornoceratids

with strongly evolute inner whorls (NYSM 12096, see Pl.32, Figs 7, 9; NYSM 12094, see Text-fig. 60F) and de-pressed whorl section; changing to subevolute outer whorlswith less depressed section and deep umbilicus at largest di-ameters known (9.1 mm, NYSM 12094). Ornament char-acterized by deep, incised grooves periodically on flanks (Pl.32, Figs 5-6) that sweep concavely forward to a slight ven-trolateral groove. On flanks, these grooves or groove-likeconstrictions increasing from approximately eight per whorlat 4 mm diameter to ten or more at 7 mm diameter, formingpart of finer ribbing pattern at larger diameters when ribscan exceed 16-18 per whorl. In middle whorls, festoon-likegrooves appearing along growth-line sinus of venter, butonly weakly developed, if at all, along ventrolateral groove;some appearing as periodic spiral grooves along ventrolateralline (NYSM 12100). Growth lines appearing to followcourse of lateral grooves, sweeping forward to slight ventro-lateral furrow, and back to U-shaped ventral sinus; lowsalient formed low on flanks. Suture showing narrowly V-shaped ventral lobe, deep, lingulate lateral lobe, and latero-umbilical saddle that is distinctly flat in its course to a lobecentered on seam (Text-fig. 60D).

Discussion.–It could be that some of the material de-scribed by Clarke under the name Tornoceras bicostatum be-longs here [Clarke, 1899a, b: pl. 8, figs 4-8; NYSM4081-4085, refigured by Miller, 1938: pl. 30, figs 5-9, as T.(Aulatornoceras) bicostatum]. This material, recorded as beingfrom near Angola and probably from the lower AngolaShale, is excluded from the account given above, which isbased on well-localized material.Attention has been drawn earlier to the possibility that

outer whorls included here under Aulatornoceras auris couldbelong to this species but this cannot be determined unlessinner whorls are present. It is clear, however, that the earlywhorl of specimens referred to A. auris at Büdesheim are dif-ferent.One specimen now included here (BMNS E 22464) was

formerly assigned to Aulatornoceras rhysum; the specimenwas said to be from the Hanover Shale at Hampton Brook(Erie County), but in view of the material collected of thisspecies at Hampton Brook (Loc. 74), it is more likely that

Table 33. Biometric data for Aulatornoceras paucistriatum (d’Archiac & de Verneil, 1842) from the lower Angola Shale.


NYSM 12094 9.1 5.1 4.8 2.6 1.06 0.29NYSM 12095 7.5 ca. 4.4 3.3 2.6 1.33 0.35NYSM 12096 4.2 ca. 2.1 ca. 1.18 1.8 1.78 0.43



the horizon was the lower Angola Shale.The horizon of the original German type material is not

known exactly; the localities of Adorf and Oberscheld pro-duce aulatornoceratids at several levels in the Frasnian(Adorfian). Most records of the species elsewhere have notincluded descriptions thorough enough to enable compari-son with the New York material.

Distribution.–Lower Angola Shale (West Falls Group;Erie and Wyoming counties).All New York specimens, except three, from either the

Point Breeze Goniatite Bed or concretionary bed immedi-ately below it. BMNS E 22464, from an uncertain horizonin Hampton Brook as noted above; NYSM 12100, Hamp-ton Brook (Erie County; Loc. 74/3); 3226/3, Sheldon Creek(Erie and Wyoming counties; Loc. 78/4b).Point Breeze Goniatite Bed or bed immediately below:

NYSM 12095-12099, 12101, and 3235/4, 9, 13, HamptonBrook (Erie County; Loc. 74/6a); 3228/1, 4, 5, 9, 3229/15,16, 25, 34, 35, 38, and 3230/4, 7, 8, 10, Cazenovia Creek(Erie CountyLoc. 75/6).

Range.–In New York: Frasnian: Neomanticoceras Geno-zone UD I-J. Regional Zone of Sphaeromanticoceras rhyn-chostomum (22b). Conodont Zone MN 11.

Aulatornoceras rhysum (Clarke, 1898)Pl. 32, Fig. 1; Text-fig. 60E

Tornoceras rhysum Clarke, 1898, 1899a, b: 121, text-fig. 100, pl.8, fig. 14.

Tornoceras (Tornoceras) rhysum.Miller, 1938: 156-157, pl. 30, fig.11.

Aulatornoceras rhysum (pars). House, 1965: 119-121, pl. 11, fig.124, text-fig. 13F.

Type material.–The syntypes were described and illus-trated by Clarke and Miller. A lectotype (NYSM 4092) wasselected by House (1965: 119) who included a specimen(BMS E 22464) in the description that is here referred toAulatornoceras paucistriatum. The type material appears tobe wholly from the lower Hanover Shale at Java (WyomingCounty; Loc. 92).

Discussion.–No new material is assigned here. The typematerial leaves much to be desired although there can be lit-tle doubt as to the generic assignment.

Distribution.–Lower Hanover Shale (West Falls Group;Wyoming County). The type material, all that is recognized,is reported from the lower Hanover Shale, Java (WyomingCounty).

Range.–Frasnian: Archoceras Genozone UD I-K-?L. Re-gional Zone of Delphiceras cataphractum (23). ConodontZone MN 12-?13.

Genus TRUYOLSOCERASMontesinos, 1987Type species.–Tornoceras sandbergeri Foord & Crick,

1897, nom. nov. pro Gon. undulatus G. & F. Sandberger(1851: 101, pl. 10, figs 17, 17a), non Brown, 1841. TheSandbergers’ specimen is from the Frasnian of Büdesheim,Germany.

Diagnosis.–Tornoceratids with nearly closed umbilicus;laterally compressed adults with ventrolateral furrows, sim-ilar to Aulatornoceras but without the festoons or constric-tions.

Included species.–Truyolsoceras sandbergeri (Foord &Crick, 1897); T. bicostatum (Hall, 1843: 246, text-fig.107(8); holotype AMNH 5888/1); and T. clarkei (Miller,1938: 80-81, pl. 14, figs 15-17; cotypes NYSM 5652-5654).

Discussion.–Separation of these forms from Aula-tornoceras without considerably more work on biostratigra-phy and evolution of the aulatornoceratids was premature.As it stands, Truyolsoceras is used for aulatornoceratids with-out the morphological extreme festoons of the aurisGroup,ribbing of the paucistriatum Group, or constrictions as inthe constrictum Group (see groups discussed under Aula-tornoceras above). Because such forms occur in the Givetianit could be the stock from which others are derived.

Truyolsoceras bicostatum (Hall, 1843)Pl. 27, Figs 7-10, Pl. 30, Figs 2, 4-14;Text-figs 59B, 60A-C, H; Table 34

Goniatites bicostatus Hall, 1843: 245-246, text-fig. 107(8).Tornoceras (Aulatornoceras) bicostatum (pars). Miller, 1938: 167-

170, pl. 14, fig. 5, pl. 32, fig. 1.Aulatornoceras bicostatum. House, 1962: 262; 1965: 120-121, pl.

10, figs 111-116, pl. 11, figs 125-137, 139-142, text-figs 16A-D.

Truyolsoceras bicostatum.House & Kirchgasser, 1993: 276; Becker,1993a: 218.

Type material.–Hall’s type material is AMNH 5888/1,lectotype here designated (Hall, 1843: 245, text-fig. 107 (8);1879: pl. 72, fig. 9), AMNH 5888/1:1 (Hall, 1879: pl. 72,fig. 8, pl. 74, fig. 1), and AMNH 5888/1:2 (Hall, 1879: pl.72, fig. 10), all from the Gowanda Shale at Corell’s Point,Lake Erie, near Brockton (Chautauqua County).

New York material.–As described by House (1965), in-cluding new collections, all from the Gowanda Shale(Canadaway Group).

Dimensions.–See Table 34 and Text-fig. 59B.Description.–A description based solely on material from

the type locality and horizon, using 23 specimens, was pub-lished by House (1965), so this species will not be re-described although illustrative material is shown herein.

Discussion.–Further statistical information in addition tothat published in 1965 is shown on the graphs for thisspecies (Text-fig. 59B); some of this material is from locali-ties for the Corell’s Point Goniatite Bed other than the typelocality at Corell’s Point. Data incorporated in the graphs(Text-fig. 59B) are based on NYSM 12121-12124, all fromthe Corell’s Point Goniatite Bed inWalnut Creek (Loc. 99b)above the railroad culvert. NYSM 12120 is from 0.9 m (3ft) above the 50-100 mm (2-4 in) siltstone bed that formsthe base for Mixer Road bridge on the northern side atForestville (Chautauqua County; Loc. 99a). NYSM 12119is from a fallen block at the same locality but thought to befrom 3 m (10 ft) above the foundation siltstone.One of the more noticeable features of this Famennian

species is the way in which the ventrolateral furrows aremore decidedly on the outer flanks than in the earlier Frasn-ian forms described here. The species differs substantiallyfrom Aulatornoceras paucistriatum of the lower Angola Shale(which was mixed with it by Clarke and Miller in their de-scriptions), especially in the much more closed-umbilicateearly whorls, the asymmetric rather than lingulate laterallobe (Text-figs 60A-C), and the absence of lateral groovesand constrictions and of ventral festoons. The species iscloser to A. aff. eifliense of this account, but the outer whorlsof those specimens are unknown, and outer whorls that areknown from the Angola and Hanover Shale specimens seemdifferent.All the material is from the Corell’s Point Goniatite Bed

apart from two specimens, referred to Truyolsoceras cf. bi-costatum, from near the road bridge north of Forestville(Loc. 99a) (NYSM 12119 and 12120). These specimenshave a much more closed umbilicus than the presumablyyounger T. clarkei from the Forestville section (compare Pl.30, Fig. 15, with Pl. 30, Figs 1, 3).

Distribution.–Corell’s Point Goniatite Bed, GowandaShale (Canadaway Group; Chautauqua and Erie counties).

Known only from the Corell’s Point Goniatite Bed, apartfrom NYSM 12119 and 12120, referred to Truyolsoceras cf.bicostatum, noted above from below that level at WalnutCreek (Chautauqua County; Loc. 99a). Noted at most lo-calities of the Corell’s Point Goniatite Bed between Corell’sPoint, Lake Erie Shore (Chautauqua County; Loc. 97), andClear Creek (North Branch), Marshfield (Erie County; Loc.106).

Range.–Famennian: Cheiloceras (Cheiloceras) GenozoneUD II-C. Regional Zone of Cheiloceras amblylobum (26).

Truyolsoceras clarkei (Miller, 1938)Pl. 30, Figs 1, 3

Gephyroceras cf. G. domanicense Clarke, 1904: 345, 380.Manticoceras clarkeiMiller, 1938: 80-81, pl. 14, figs 15-17.Aulatornoceras clarkei. House, 1962: 262; 1965: 121-124, pl. 11,

figs 122-123.Truyolsoceras clarkei. House & Kirchgasser, 1993: 276.

Type material.–The cotypes (NYSM 5652-5654) weredescribed in the works cited above and two of the specimensare reillustrated here (Pl. 30, Figs 1, 3).

Discussion.–This species is like Truyolsoceras bicostatumbut differs in a wider umbilicus and strongly marked peri-odic ribbing on the flanks. The first impression that thesemight be that species in a different preservation is notthought to be so, and material in a similar preservation lowerin the Gowanda Shale than the Corell’s Point Goniatite Bedat Forestville is here described as T. cf. bicostatum.

Distribution:–Gowanda Shale (Canadaway Group;Chautauqua County). Described only from Walnut Creek,Forestville (Chautauqua County; Loc. 99), but the strata-graphic level is not known. House (1965) suggested that itmight be from the upper Gowanda Shale, Laona Siltstone,orWestfield Shale, but these levels have not been adequately

Table 34. Biometric data for Truyolsoceras bicostatum (Hall, 1843) from the lower Gowanda Shale. See Text-fig. 59B for graph.


AMNH 5888/1 22.5 — ca. 11.3 — — —(lectotype)

AMNH 5888/1:1 ca. 25.0 10.5 12.3 2.8 0.85 0.11NYSM 6679/B 21.1 8.8 10.5 2.9 0.84 0.14NYSM 6629C 18.2 8.7 8.5 1.9 1.02 0.10USNM 137667 ca. 18.0 — ca. 8.8 — — —NYSM 11958 15.3 7.5 8.5 1.9 0.88 0.12NYSM 6679A 13.7 6.7 7.4 ca. 1.7 0.91 0.12AMNH 5888/1:2 7.3 — 3.1 1.8 — 0.25NYSM 11957 6.2 3.2 3.2 — 1.00 —NYSM 11962 5.0 2.8 3.1 — 0.90 —NYSM 11960 1.3 0.85 0.6 0.34 1.42 0.26NYSM 11962 1.12 0.93 0.62 — 1.50 —


searched in this study.Range.–Famennian: Cheiloceras (Cheiloceras) Genozone

UD II-C. Regional Zone of Truyolsoceras clarkei (25).

Superfamily CHEILOCERATOIDEA Frech, 1897(= DIMEROCERATACEAE nom. trans. Bartzsch &

Weyer, 1988)Family CHEILOCERATIDAE Frech, 1897Genus CHEILOCERAS Frech, 1897

Type species.–Goniatites subpartitus Münster, 1839, bysubsequent designation (Wedekind, 1918: 144).

Diagnosis.–Cheiloceratids with conch in earliest stagesinvolute to subevolute, in later stages with closed umbilicus.Usually rotund in early stages, later stages can be rotund tolaterally compressed with venter round to oxyconic; rarelywith gentle ribs, commonly with internal shell varices form-ing constrictions on internal mold. Basic stocks with convexgrowth lines. Suture EALI to EALU1I.

Discussion.–The question of the name of this group isimportant. No international rules govern taxa above thefamily level below which adherance to priority is required.This is true even of the Code (ICZN, 1999, effective fromJanuary 1, 2000). The principle reason for this is to avoid re-placement of well-known and understood names by the va-garies of priority. Frech (1897) established a name group forthe important genus Cheiloceras that became the best-knownFamennian goniatite whenWedekind (1918) used it for theCheiloceras Stufe. The strength of Wedekind’s views are em-phasized by his suborder Cheiloceracea. Proposals to replacethe current group name used here by Dimeroceratacea orDimeroceratoidea are misconceived and misjudge the im-portance of stability in high-taxon terminology, which is thebasis of exclusion from priority rulings of all recent Codes.In his detailed review of Nehdenian faunas, Becker

(1993a) subdivided this genus into four subgenera: Cheilo-ceras s. s., Raymondiceras, Staffites, and Puncticeras. Only thefirst is represented in New York. Strand (1929: 8) wrongly

stated that Cheiloceras Trouessart, 1898, had priority, andproposed as replacements the names Cheilocerotes andCheiloceratos, both of which are invalid (see Richter, 1929:38); there is also ChilocerasDreverman (1901: 123), a nom.van. As has been elegantly shown by Becker (1993b: 128),this genus plays a major role in the radiation of goniatitesfollowing the Upper Kellwasser Event (Text-figs 20-21). Therecognition of the genus in North America was relativelylate, first in New York, and then in northwestern Canada(House, 1962; House & Pedder, 1963).

Distribution.–Worldwide except South America andAntarctica. Marker genus for early Famennian.

Range.–Famennian: UD II-B to UD III-C; if Hember-gian Raymondiceras are included, UD IV-A; otherwise ex-tinct in UD II-G.

Cheiloceras (Cheiloceras) amblylobum(G. & F. Sandberger, 1851)

Pl. 35, Figs 1-10; Text-figs 59C, 61A-B; Table 35

Goniatites retrorsus var. amblyloba G. & F. Sandberger, 1851: 108,pl. 10a, figs 20, 24, pl. 10b, fig. 6; 1852: 108.

Cheiloceras amblylobus.Wedekind, 1918: 146.Cheiloceras (Cheiloceras) amblylobum. House, 1962: 274-276;

Becker, 1993a: 244-246, text-figs 81b, 82e, 83f, pl. 14, figs10-11; Kirchgasser & House, 1981: 45-46, 49; House &Kirchgasser 1993: 276.

Type material.–On the assumption that the originals ofthe Sandbergers are lost, Matern (1931b) designated a neo-type; but House, leaving open the question of loss, chose aslectotype the specimen favored by Wedekind (1918: 146)and figured by the Sandbergers (1852: pl. 10, fig. 8). Becker(1993a: 245, text-fig. 83f, pl. 14, figs 10-11) has designateda new neotype as Museum für Naturkunde (Berlin), Be1256, from the Nehden-Schurbusch, Rhenish Slate Moun-tains.

NewYork material.–Fifteen pyritic molds were listed pre-

Table 35. Biometric data for Cheiloceras (Cheiloceras) amblylobum (G. & F. Sandberger, 1851) from the lower Gowanda Shale. See Text-fig. 59C for graph.


NYSM 16552 22.7 13.2 13.9 0 0.95 —USNM 137665 14.6 9.1 7.1 ca. 1.0 1.28 0.07USNM 137666 13.7 ca. 9.0 7.8 ca. 1.0 1.15 0.07NYSM 16550 13.4 9.3 7.9 ca. 0.5 1.18 0.04NYSM 16557 12.8 8.9 7.2 ca. 0 1.24 —NYSM 16554 10.4 7.4 7.2 0 1.03 —NYSM 16551 9.7 7.3 5.2 ca. 0.5 1.40 0.05NYSM 16553 8.5 6.2 4.8 ca. 0 1.29 —NYSM 16556 6.9 5.9 3.8 0.5 1.55 0.07NYSM 16555 3.8 2.7 2.4 0 1.12 —


viously (House, 1962: 276), including USNM 137665 and137666 collected by W. Moran, NYSM 11239-11242 col-lected by J. M. Clarke and D. D. Luther in 1898, and spec-

imens in the collections of MRH now deposited with theNYSM with other material collected at several localities.

Dimensions.–See Table 35 and Text-fig. 59C.Description.–A description of material from Corell’s

Point was given earlier (House, 1962: 274-275) and neednot be repeated. Statistical details of additional material isincluded in Table 35 and Text-fig. 59C. Typical sutures areillustrated in Text-figs 61A-B, and a range of specimens areillustrated on Pl. 35.

Distribution.–In New York: Corell’s Point Goniatite Bed,Gowanda Shale (Canadaway Group; Chautauqua, Catta-raugus, Erie, and Wyoming counties).From Chautauqua County: Lake Erie Shore at Corell’s

Point (Loc. 97), Little Canadaway Creek (Loc. 98), WalnutCreek (Loc. 99), and Smith Mills (Loc. 100). From Catta-raugus County: Big Indian Creek (Loc. 101), Little IndianCreek (Loc. 102), Cattaraugus Creek (Cattaraugus IndianReservation) (Loc. 103), and Cattaraugus Creek (SouthBranch) (Loc. 104). From Erie County: Clear Creek (NorthBranch), Taylor Hollow (Loc. 105), and Anthony Gulf (Loc.107). Also, there is a specimen in the NYSM from Java(Wyoming County; Loc. 109), but our work did not locatethe level.

Range.–Famennian: Cheiloceras (Cheiloceras) GenozoneUD II-C. Regional Zone of C. amblylobum (25).

Superfamily SPORADOCERATOIDEA nom. trans.Miller & Furnish, 1957

Family SPORADOCERATIDAE nom. trans.Miller & Furnish, 1957

If this group is assigned to superfamily rank, then it shouldbe referred to as Sporadoceratoidea (nom. trans. Miller &Furnish, 1957) rather than Praeglyphioceratoidea(Ruzhencev, 1957), as in Becker (1993a), to conform withthe ICZN guidance concerning the least disruption of seniortaxon names. The group comprises forms with convexgrowth lines and sutures that differ from the Cheiloceratidaein the addition of extra adventitious ventral lobes (A2 and insome A3).

GenusMAENECERAS Hyatt, 1884Type species.–Goniatites acutolateralisG. & F. Sandberger,

1850, by original designation (Hyatt, 1884: 321).Diagnosis.–Sporadoceratids with discoidal to subglobular

conchs with closed umbilicus and convex growth lines. Su-ture EA2A1LUI but with A1 sharp or narrowly rounded andA2 shallow and rounded. For discussion and includedspecies, see Becker (1993a).

Distribution.–Widespread. North America (very rare),Europe, Russia, Asia, North Africa, Australia, and SouthAmerica.

Text-fig. 61. Sutures of Cheloceras, Maeneceras, and Sporadocerasspp. from New York, Pennsylvania, and Ohio (after House, 1962,and House et al., 1986). A-B. C. (Cheiloceras) amblylobum (G. &F. Sandberger, 1851). A. USNM 137665, suture based on speci-men collected by W. Moran from the Gowanda Shale at Corell'sPoint (Loc. 97), on the shore of Lake Erie, Chautauqua County,suture reversed, X 3. B. USNM 137666, suture based on specimenfrom same locality as A, X 2.8. C-D.M. aff. acutolaterale (G. & F.Sandberger, 1850), USNM 137645, based on a specimen collectedby H. S. Williams in 1884 from the Ellicott Shale at Porter'sCreek, Summerdale, Chautauqua County. C. Suture at 83 mm di-ameter, X 0.7. D. Reversed suture at 117 mm diameter, a com-posite diagram, X 0.7. E. M. milleri (Flower & Caster, 1935),diagram of the suture (reversed), from Miller (1938) of the holo-type from Howard Quarry, Erie County, Pennsylvania, X 0.7. F.M. inflexum (Wedekind, 1908), NMNH 240513, suture based onpyritic fragments from 2 m above the base of the Cleveland Shale,Birmingham, Ohio, X 2.5.


Range.–Famennian: UD II-G to V-B (Becker, 1993b).

Maeneceras aff. acutolaterale (G. & F. Sandberger, 1850)Pl. 35, Figs 11, 13; Text-figs 61C-D, 62

aff. Goniatites acutolateraleG. & F. Sandberger, 1850: pl. 6, figs 1,1a-b; 1851: 98.

aff.Maeneceras acuto-laterale. Hyatt, 1884: 321.Sporadoceras cf. pompeckjiWedekind, 1918. House 1962: 276, pl.

46, fig. 12, text-fig. 12A.Maeneceras cf. pompeckji. Kirchgasser & House, 1981: 46, 49;

House & Kirchgasser, 1993: 276.

Type material.–Holotype ofMaeneceras acutolaterale de-scribed by the Sandberger brothers, refigured by Becker(1993a: 313, fig. 96).

New York material.–One specimen only (USNM137645) collected by H. S. Williams in 1884 from PorterCreek, Summerdale (Chautauqua County).

Dimensions.–USNM 137645, maximum D = 132 mm,D = 120 mm, WW = 69 mm, and UW = ?0 mm.

Description.–Features of shell as in Text-fig. 62. Bodychamber occupying last one-third whorl; phragmoconeshowing sutural approximation. Laterally compressed anddiscoidal with rounded venter and closed umbilicus. A2 lobeshowing increasing depth in last half whorl of phragmocone.Growth lines not seen.

Discussion.–At present, the distinction betweenMaeneceras and Sporadoceras lies in the depth of the A2 lobe.Yet diagnoses do not state the percentage of depth required,nor the shell diameter at which this must be attained. The

Sporadoceratidae are in need of much biometric work.Originally this species was referred to Sporadoceras cf.

pompeckji, but the types of that species are relatively smalland the extent to which the A2 lobe would deepen in laterstages is uncertain. The specimen can be compared to theholotype of Maeneceras acutolaterale, from which it differsin the more parallel-sided conch and in the lesser depth ofthe A2 lobe, but the A1 lobe is not so sharp.

Distribution.–Ellicott (Chadakoin) Shale (ConneautGroup; Chautauqua County). The specimen is labeledHSW coll. 518B. J. W. Wells informed MRH that locality518B is marked on manuscript maps of H. S. Williams,then at Cornell University, which showed it to be alongPorter Creek, 2 mi (3.6 km) northwest of Summerdale(Chautauqua County), at a point just below Porter Ceme-tery at an altitude of approximately 1,450 ft (442 m) (Loc.110 of this report), This point lies in the middle part of theoutcrop of the Ellicott (Chadakoin) Shale on the map ofTesmer (1954: 29).

Range.–In New York: Famennian:MaenecerasGenozoneUD II-G. Regional Zone of M. aff. acutolaterale (27). InGermany, the type species is said to range from ?upperbiferum Zone (UD II-G) to the contiguum Zone (UD II-H)(Becker, 1993a: 314). In Pennsylvania, the occurrence ofM.milleri (illustrated here, Text-fig. 61E) higher in the NorthAmerican succession, in the Conewango Group (Cattarau-gus-Oswayo equivalents), Howard Quarries ( Loc. 111), Re-gional Fauna 28, correlates with UD II-H (Becker &House,2000: 134). Still higher, in Ohio, M. inflexum (Text-fig.61F) from the Cleveland Shale (Regional Fauna 29), corre-lates with PrionocerasGenozone UD IV-A (Becker &House,2000).

Family and Genus Indeterminable

?Genus edwinhalli (Clarke, 1898)Pl. 35, Fig. 12

Tornoceras Edwin-halli Clarke, 1898, 1899a, b: 111, text-fig. 85.Tornoceras (Tornoceras) edwinhalli. Miller, 1938: 151, pl. 31, figs

10-11.

Discussion:–This single specimen (NYSM 4090) de-scribed by both Clarke and Miller needs no further descrip-tion. However, it seems that the drawn figure is not a ventralsuture at all, but the impression of the dorsal suture from awhorl now lost. Such simple dorsal sutures are found insome prolobitids, but in the absence of further information,and perhaps the dissection of the specimen, no more precisenaming is possible.

Distribution.–Conneaut Group, Nile (Allegany County).Range.–Famennian.

Text-fig. 62. Maeneceras aff. acutolaterale (G. & F. Sandberger,1850), USNM 137645, collected by H. S.Williams in 1884 fromthe Ellicott Shale at Porter's Creek, Summerdale, ChautauquaCounty. Scale = 10 cm.



Text-fig. 63 (at left and above). Diagramatic illustrations of zone-defining goniatites in the late Devonian ofNew York State. Units and Zones as in Text-fig. 24; at the top, read Oswayo (for Osweyo). 12. Tornoceras cf.uniangulare (Text-fig. 54D); 13. Pharciceras amplexum (Text-figs 26E, H); 15a. Epitornoceras cf. mithracoides(Text-fig. 57F), E. cf. peracutum (Text-fig. 57D); 15b. Ponticeras perlatum (Text-figs 28C-D); 15c. Chutocerasnundaium (Text-figs 28I-J); 16a. Koenenites styliophilus styliophilus (Text-figs 34F, L); 16b. K. styliophilus kilfoylein. ssp. (Text-fig. 33N); 17a. Manticoceras sinuosum apprimatum (Text-fig. 42F); 17b. K. beckeri n. sp. (Text-fig. 33A); 18. Sandbergeroceras syngonum (Text-fig. 27H); 19. Probeloceras lutheri (Text-fig. 50F); 20. Prochoritesalveolatus (Text-figs 31C-D); 21a. Naplesites iynx (Text-fig. 52A); 21b. Wellsites tynani (Text-fig. 27A); 21c.Schindewolfoceras chemungense (Text-fig. 27I); 22a. Playfordites cf. tripartitus (Text-fig. 42A); 22b. Sphaero-manticoceras rhynchostomum (Text-figs 43F, 42H); 23.Delphiceras cataphractum (Text-fig. 41G); 24a. Crickiteslindneri (Text-figs 49A-B); 24b. Sphaeromanticoceras rickardi (Text-fig. 48C); 24c1. ?Archoceras sp. (Text-fig.48G); 24c2. ?Crickites sp. juv. (Text-fig. 48H); 25. Cheiloceras amblylobum (Text-fig. 61A); 27.Maeneceras aff.acutolaterale (Text-fig. 61C); 28.Maeneceras milleri (Text-fig. 61E). At top left, read Oswayo.


SYNOPSISText-fig. 63 summarizes the succession of zone-defining go-niatites in the late Devonian of New York State with illus-trations of sutures and whorl-sections of some key taxa.

ADDENDUMNoted here are recent reports concerning some of the taxaand sections described in this monograph. Work et al.(2007) reported the discovery of Pharciceras in the New Al-bany Shale (Trousdale Member) in Kentucky. Pharcicerasbarnetti Work et al., 2007, is similar to P. amplexum fromthe Upper Tully Limestone but differs in having a widerconch and more depressed whorl form. The two horizonswith P. barnetti correlate to an interval within the lowerGeneseo Shale of the Taghanic Onlap sequence that spansthe level of the Fir Tree Limestone (and equivalent LeicesterPyrite) of the upper Givetian (MD III A; hermanni and dis-paralis conodont zones).Baird et al. (2006a) have greatly refined the sections of

the lower to middle Genesee Group between Lake Erie andthe Genesee Valley. The delineation of the many discontinu-ities, and particularly those associated with the Lodi andLinden Horizons and the Lower and Upper Divisions of theGenundewa Limestone, should lead to an even finer-scaleresolution of the goniatite succession.Zambito et al. (2007, unpublished data) have revised

and analyzed the Sherburne, Renwick, and Ithaca Forma-tions and equivalents of the Genundewa Limestone andWest River Shale in the Ithaca area using the methodologyof sequence stratigraphy. Newly measured sections includethe classic section at Fall Creek, Ithaca. The goniatite succes-sion outlined herein is illustrated in a composite section ofthe lower, middle, and part of the upper Genesee Group.Several submember and system-tract units are tentativelycorrelated with the Genesee Group succession to the west.The new sequence stratigraphy provides a framework for re-fining the ranges of genera and species of goniatites in theIthaca region.Finally, Baird et al. (2006b) reported the discovery of

Naplesites in the lower Rhinestreet Shale at Lake Erie shore.The occurrence is a bedding plane surface in black shalewith numerous flattened specimens (some partially pyritizedand showing sutures) and a few uncrushed specimens inscattered concretions at the same level. The species is likelyto be eitherN. inyx orN. naplesense, the species described byClarke (1898) from an unknown level around Naples, On-tario County. The discovery of Naplesites at Lake Erie con-firms our guess that Clarke’s specimens came from the lowerRhinestreet Shale. Work has begun to see if the Naplesiteshorizon at Lake Erie can be located in the section at Naples.-WTK

ACKNOWLEDGMENTSProgress on this work has proceeded over many years andconsequently we are indebted to many for their help and as-sistance. The Natural Environment Research Council(NERC) in the 1960s funded a research project to MRHthat enabled us both to commence a systematic study of theLate Devonian across the State of New York during whichtime stratigraphical successions were established in a generalway and systematic collections begun. The New York StateMuseum and Science Service funded field work at that timeby WTK and provided a vehicle for two seasons’ work and,through the kind offices of its staff, helped in many ways.We thank especially L. V. Rickard for putting his enormousknowledge of the Devonian geology of the State at our dis-posal; D. W. Fisher, former State Paleontologist, who facil-itated work in every way; and C. Kilfoyle for guidance andloans from the outstanding State collections at Albany, andfor permission to photograph and access the State collec-tions. More recently, E. Landing and L. Van Aller Hernackhave provided access to the collections and C. A. VerStraeten has given bibliographic help. At the University ofRochester, R. G. Sutton graciously provided data on blackshale correlations that proved to be crucial in determiningthe Rhinestreet equivalent goniatite sequences, and G. Klochas generously shown us his collections. Jean Dougherty ofthe Geological Survey of Canada has advised on the materialcollected by E. M. Kindle.To Cornell University, and the late Professor J. W.Wells

in particular, we owe a debt that cannot be repaid in words.John Wells introduced both of us to the geology of centralNew York and supervised the doctorate by WTK on theCashaqua Shale. Throughout the period until his untimelydeath, he gave his whole-hearted encouragement and puthis encylopedic knowledge and substantial collections of fos-sils, books, and maps at our disposal. With his marvellouswife, Pie Wells, he provided respite and refreshment count-less times during fieldwork and odd visits to their home,Lucky Stone Lodge, at Sheldrake on the banks of CayugaLake.The Paleontological Research Institution (PRI) at Ithaca

has provided material for study, as has the Buffalo Museum.There was a major early input by the U. S. National Mu-seum when G. A. Cooper introducedMRH to many aspectsof North American Devonian work, and material in the Mu-seum collections has been made available for study. The helpof others at the Museum, especially W. A. Oliver, Jr., HelenDuncan, Jean Berdan, and J. T. Dutro, Jr., is much appre-ciated. At the American Museum of Natural History, NewYork, Otto Haas and N. H. Landman have facilitated accessto collections, as have N. Nitecki and S. Lidgard at the Field


Museum of Natural History in Chicago. At the PRI, W. D.Allmon andW.Taylor have facilitated our work. At the Uni-versity of Iowa, B. F. Glenister and W. M. Furnish havemade Devonian comparative material available and, over thelast decade, collaboration with the conodont biostrati-graphic studies of G. Klapper has sharpened aspects of ourwork. More recently we have appreciated demonstration oftheir work on Devonian rocks in New York by C. E. Brett,G. Baird, and D. J. Over. We are both greatly indebted forastute comments and help by our colleague, R. T. Becker, ofthe University of Münster, Germany, for his collaboration infieldwork in New York as well as in Australia, Russia, Eu-rope, and North Africa, and for his systematic collation ofliterature that he has put freely at our disposal.This work has continued while MRH was successively a

member of Geology Departments at Oxford, Hull, andSouthampton Universities, where staff provided secretarialand photographic help; J. Garner and B. A. Marsh providedmost of the prints used on the plates. Many thanks are dueto Felicity House for continuing support.At the State University of New York at Potsdam, WTK

has been supported since 1969 by numerous grants fromthe SUNY Research Foundation, the New York State UnitedUniversity Professions, and Potsdam College. The relatedcollaboration on conodonts with G. Klapper at the Univer-sity of Iowa was supported by National Science FoundationResearch Opportunity Award (EAR-890475) to WTK.Thanks also are due to the many undergraduate studentswho assisted WTK on the conodont work; they are ac-knowledged by name in separate publications. The generoussupport over the years of many colleagues at Potsdam Col-lege is greatly appreciated, particularly the staffs of the Officeof Scholarships and Grants, Crumb Library, and ComputerServices; special thanks go also to R. Bitely for his adviceand help on photographic work.We are also indebted to ourfriends on the faculty and staff of the SUNY Potsdam Ge-ology Department, Professors F. Revetta, B. Van Diver, N.O’Brien, J. Carl, and R. Badger, J. Chiarenzelli, L. Amatiand the Departmental Secretaries, J. Moriarty, V. O’Brien,and J. Bullis, for their years of support and encouragement.Thanks especially to J. Bullis, for preparing the final type-script, and Patty Stone for much technical assistance.Thanks also to Kim Coleman for computer-drafting Text-figs 1 and 63.We greatly appreciate the timely reviews of the manu-

script provided by Gordon Baird (SUNY Fredonia), BrianGlenister (University of Iowa), and Warren Allmon (Pale-ontological Research Institution); most of their suggestionshave been incorporated in this final version. A few days afterreceiving the reviews and notification of acceptance for pub-lication of the manuscript, Michael House passed away (06

August 2002) in Dorchester near his home in Weymouth,Dorset, England. The task of editing this final manuscriptthus fell to me (WTK). Some parts needed updating but Ihave made every effort to keep the content and style as closeas possible to the original. Thanks to my new colleagues atthe Department of Geology for supporting the project invarious ways: M. Rygel, C. Kelson, and R. Greene (Secre-tary). Thanks also to Betsy Northrop for twenty-fve years ofsupport and patience. Finally, special thanks to Nancy Dutrofor a fine job of copy-editing for PRI and to PaulaMikkelsen, Director of Publications at PRI, for the final ed-iting and putting the parts together for publication. All er-rors remaining are of course my responsibility.Michael House and I agreed long ago to dedicate this

monograph to the memory of Professor andMrs. JohnWestWells.

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PLATES

204 BULLETINS OF AMERICAN PALEONTOLOGY no. 374BULLETINS OF AMERICAN PALEONTOLOGY no. 374204


PLATE 1

Figure Page1–8. Pharciceras amplexum (Hall, 1886) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 90

1-2. USNM 96545a, lateral and ventral views of a specimen from the West Brook Member of the Tully Limestone, 1mi (1.6 km) northeast of Laurens, Otsego County (Loc. 9b), X 0.8.

3. NYSM 3729, lateral view of the holotype figured by Hall (1886: pl. 127, fig. 1) from the Tully Limestone nearLodi Landing, Seneca Lake, Seneca County (Loc. 3), X 1.6 (reversed for comparison).

4. NYSM 11999, lateral view of a specimen from the Pharciceras Bed, West Brook Member of the Tully Limestone,June's Quarry, Tully, Onondaga County (Loc. 7), X 1.2.

5-6. USNM 143014, lateral and ventral views of a specimen found loose in June's Quarry, Tully, Onondaga County(Loc. 7) by W. A. Oliver, Jr., X 1.2.

7. NYSM 12002, lateral view of a specimen showing fine ribbing on the early whorls from the Pharciceras Bed, WestBrook Member of the Tully Limestone, June's Quarry, Tully, Onondaga County (Loc. 7), X 1.6.

8. USNM 96545b, lateral view of a specimen showing the development of a double ventrolateral furrow from theWest Brook Member of the Tully Limestone, 1 mi (1.6 km) northeast of Laurens, Otsego County (Loc. 9b), X0.8.

9. Wellsites tynani House & Kirchgasser, 1993 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 96SUI 42418, holotype from the Moreland Shale (basal tongue of the Rhinestreet Shale) found in spoil from a roadcutfor US Rte. 17 at Elmira, Chemung County (Loc. 70), X 0.8. Photograph by M. C. Tynan.

10. Pharciceras tridens (G. & F. Sandberger, 1849) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 90Wiesbaden Museum [unnumbered], ventral view of the lectotype, here designated, a specimen figured by G. & F.Sandberger (1849: pl. 4, fig. 2) from Oberscheld (Königzug), Germany, X 1.2.

PLATE 2

Figure Page1-2. Sandbergeroceras? enfieldense n. sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 95

PRI 8742, latex mold of a specimen from equivalents of the Cashaqua Shale at the top of Bald Hill, near Brookton-dale, Tompkins County (Loc. 94), coll. S. Hollister. 1, X 0.8; 2, X 1.6.

3. ?Schindewolfoceras sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 96NYSM 12025, lateral view of a specimen from the "Chemung" between Chemung and North Chemung, ChemungCounty (Loc. 95), X 0.8.

4-7. Sandbergeroceras syngonum (Clarke, 1897) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 934-5. NYSM 4076, lateral and ventral views of a gutta percha mold of a specimen from a concretion, probably from

the Cashaqua Shale, near Naples, Ontario County, figured by Clarke (1898: pl. 7, fig. 19), X 1.3.6. NYSM 4075, lateral view of a baritic replacement from "crinoidal blocks," probably from the Cashaqua Shale, at

Naples, Ontario County, figured by Clarke (1898: pl. 6, fig. 23), X 4. Photograph by J. W. Wells.7. NYSM 12026, lateral view of a specimen collected by H. S. Williams from the upper Middlesex Shale equivalent,

near Harford Mills, Tioga County (Loc. 64), X 1.6. See also Pl. 3, Fig. 6.

8. Schindewolfoceras chemungense (Vanuxem, 1842) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 97NYSM 4073, lateral view of the holotype figured by Vanuxem (1842: text-fig. 49, fig. 1) probably from the CayutaShale (Rhinestreet Shale equivalent), "near Owego," Tioga County (Loc. 69), X 1.6.




PLATE 3

Figure Page1. Schindewolfoceras? aff. equicostatum (Hall, 1874) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 98

NYSM 12027, latex mold of a specimen from the "Upper Cayuta Chesney Formation" (middle Rhinestreet Shaleequivalent) collected by J. W. Wells from a small quarry alongside States Land Road in Fairfield State Forest, 0.4 mi(0.64 km) east of Fleet Road, 1.5 mi (2.4 km) southwest of Speedsville, Tioga County (Loc. 67), X 0.8.

2-3. Sandbergeroceras? enfieldense n. sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 95NYSM 12028, latex mold of the holotype collected by K. Caster from the “Enfield” (Cashaqua equivalent) at Uni-versity Quarries (Loc. 66), southeast of Ithaca, Tompkins County. 2, X 0.8; 3, X 2.4.

4. Schindewolfoceras? equicostatum (Hall, 1874) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 97NYSM 4074, lateral view of the holotype figured by Hall (1876: pl. 69, fig. 10) from a boulder near Athens, Penn-sylvania, probably from a level equivalent to the lower West Falls group, X 1.1. Based on a photograph of a cast.

5-6. Sandbergeroceras syngonum (Clarke, 1897) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 935. NYSM 4077, lectotype, here designated, of a syntype figured by Clarke (1898: pl. 7, fig. 20) from the Middlesex

Shale at Snyder's Gully, near Naples, Ontario County (Loc. 46c), X 0.8.6. NYSM 12026, latex cast of a specimen collected ca. 1905 by H. S. Williams (his loc. Hd 129/1b) from between

1,200 and 1,300 feet (365-396 m) on the western side of the valley (said to be from the Enfield Shale; probablyupper Middlesex Shale equivalent), 1.5 mi (2.4 km) south of Harford Mills (Loc. 64), X 0.8. Photograph pro-vided by J. W. Wells.


PLATE 4

Figure Page1-4. Schindewolfoceras aff. chemungense (Vanuxem, 1842) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 97

SUI 40080, specimen collected by J. Regan from a roadcut on Hwy 19 near Big Spring, Missouri, collected as floaton a Calloway Limestone exposure, but B. F. Glenister informs us that it is probably from the Burlington Limestone,X 0.8.

5-9. Ponticeras perlatum (Hall, 1874) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 995. Photograph provided by J. W. Wells of specimens in original vertical and horizontal orientations in a concretion

from the base of the Sherburne Formation (Lodi Limestone), in Romulus Town Quarry, 1.5 mi (2.4 km) north-west of Ovid, Seneca County (Loc. 27a), X ca. 0.4.

6. CU 40161, specimen collected by Mrs. Grealey from the Upper Geneseo Shale, Taughannock Gorge, TompkinsCounty, X 0.8. Photograph provided by J. W. Wells.

7. NYSM 13082, enlargement of the inner whorls of the specimen figured on Pl. 5, Figs 1-4 from the lower part ofthe Sherburne Siltstone (Lodi Limestone) in Lodi Glen, Seneca Lake (Loc. 27), X 17.8.

8. NYSM 5122, specimen mentioned by Clarke (1898: 86), probably from the Sherburne Siltstone, Ithaca, TompkinsCounty, X 1.2.

9. NYSM 3649, lectotype figured by Hall (1876: pl. 70, fig. 12; 1879: pl. 71, fig. 12) and mentioned by Clarke(1898: 86) from the Sherburne Siltstone, Homer, Cortland County, X 1.1.




PLATE 5

Figure Page1-9. Ponticeras perlatum (Hall, 1874) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 99

1-4. NYSM 13082, specimen from the lower part of the Sherburne Siltstone (Lodi Limestone) in Lodi Glen, SenecaLake (Loc. 27), collected by J. W. Wells, X 2.4. See also Pl. 4, Fig. 7.

5-6. NYSM 13084, specimen from the Lower Sherburne Siltstone (Lodi Limestone), Sheldrake Springs, SenecaCounty, J. W. Wells coll. 40118 (Loc. 308). 5, X 1; 6, X 2.2.

7-9. NYSM 13083, specimen from the lower Sherburne Siltstone (Lodi Limestone) at Lodi Glen (Loc. 27), nearLodi, Seneca County, J. W. Wells Coll. 400117. 7, X 0.8; 8-9, X 1.2.


PLATE 6

Figure Page1-15. Acanthoclymenia neapolitana (Clarke, 1892) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 103

All specimens are baritic shell replacements apparently from the level of the Shurtleff Septarian Horizon in theCashaqua Shale, probably from Shurtleff ’s Gully (Loc. 41), Livingston County, or immediately adjacent areas.1. NYSM 3629, lateral view of a cotype figured by Clarke (1892: 63, text-fig. 11; 1898: pl. 8, fig. 25), X 3.2.2-3, 10, 14. NYSM 3625, various views of the lectotype, a cotype figured by Clarke (1892: 63, text-fig. 3). 2-3, X ca.

3.2; 10, 14, X ca. 5.4-5. NYSM 11264, lateral and ventral views of early stages based on a specimen among Clarke's material, X 5.6.6-8. NYSM 3634, apertural, lateral, and ventral views of a specimen figured by Clarke (1898: pl. 8, fig. 24), X 6.9. NYSM 3633, lateral view of a specimen figured by Clarke (1898: pl. 8, fig. 22), X 4.4.11. NYSM 3628, lateral view of a specimen figured by Clarke (1898: pl. 8, fig. 21), X 5.12. NYSM 3632, lateral view of a specimen figured by Clarke (1898: pl. 8, fig. 20), X 4.4.13. NYSM 3634, lateral view of a specimen figured by Clarke (1898: pl. 8, fig. 19), X 2.4.15. NYSM 3624, lateral view of a specimen among Clarke's material, X 5.




PLATE 7

Figure Page1. Koenenites sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 112

NYSM/CU 40022, lateral view of a specimen figured by Wells (1956a) from the West River Shale, 0.6 mi (0.96 km)south of Hicks Point (Loc. 29a), western side of Canandaigua Lake, Ontario County, X 0.8.

2-3. Neomanticoceras paradoxum (Matern, 1931) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 160SM XI 320c, whorl cross section and lateral views of a specimen collected by MRH from the Frasnian BüdesheimerSchiefer about 150 m southwest of the church at Büdesheim, Eifel, West Germany, X 6.

4-6. Prochorites alveolatus (Glenister, 1958) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 109NYSM 4063, enlarged views showing the wrinkle layer and ventral and lateral views of a specimen figured by Clarke(1898: pl. 7, fig. 4) probably from the Shurtleff Septarian Horizon, Cashaqua Shale, near Honeoye Lake. 4, X 8; 5-6,X 4.


PLATE 8

Figure Page1-4, 6, 9. Koenenites styliophilus styliophilus (Clarke, 1898) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 113

1-2. NYSM 3783, lectotype, here designated, figured by Clarke (1898: pl. 6, fig. 30; text-fig. 55), the only survivingsyntype; recorded by Clarke as from the Naples Beds, Naples, Ontario County; probably from the upper PennYan Shale, X 1.2.

3. NYSM 12134, lateral view showing growth lines and ventrolateral furrow, Upper Penn Yan Shale, Linden GoniatiteHorizon, Linden Falls (Loc. 15a/8), Linden, Genesee County, X 1.

4. CU 40101, lateral view of a specimen collected by J. W. Wells from the upper Penn Yan Shale about 3 m (10 ft)below the top layer of the Genundewa Limestone and probably from the Linden Horizon, Taunton Gully (Loc.17/6), near Leicester, Livingston County, X 0.8.

6. NYSM 12135, lateral view of a specimen from the Crosby Sandstone, Sunset Point Gully (Loc. F, Py-16/1), KeukaLake, 9.3 km (5.8 mi) southwest of Penn Yan, Yates County, X 0.8.

9. NYSM 16563, lateral view of a specimen collected by D. D. Luther (Loc. 3397), Crosby Sandstone at top of PennYan Shale, Sartwell Ravine (Loc. C, Py-11), 1.6 km (1 mi) south of Penn Yan, Yates County, X 0.8.

5, 7-8, 12-13. Koenenites styliophilus kilfoylei n. ssp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1195. NYSM 16558, lateral view of a specimen from the Genundewa Limestone exposed along US Rte. 20, Bethany

Center (Loc. 24), Genesee County, X 1.2.7-8. NYSM 12143, lateral and ventral views of the holotype from the Genundewa Limestone exposed along US Rte.

20, Bethany Center (Loc. 24), Genesee County, X 1.2.12-13. NYSM 12144, lateral and ventral views of a specimen (morphotype or form A) from the Genundewa Lime-

stone exposed along US Rte. 20, Bethany Center (Loc. 24), Genesee County. 12, X 1.6; 13, X 1.2.

10-11, 14. Koenenites beckeri n. sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12110-11. NYSM 12149, lateral and whorl-section views of a specimen from a loose concretion found 3 m (10 ft) below

the Bluff Point Siltstone, West River Shale, in the gully 1.3 km (0.8 mi) south of Middlesex (Loc. XX, Nap-8),Yates County, X 1.6.

14. NYSM 12150, lateral view of the holotype from a loose concretion found 3 m (10 ft) below the Bluff Point Silt-stone, West River Shale, in the gully 1.3 km (0.8 mi) south of Middlesex (Loc. XX, Nap-8), Yates County, X 1.2.




PLATE 9

Figure Page1. Chutoceras nundaium (Hall, 1874) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 107

NYSM 3753, lateral view of a specimen figured by Hall (1888: pl. 128, fig. 1) thought to be from the Ithaca Forma-tion, Genesee Group, University Quarry, Fall Creek (Loc. X, Dy-10), Ithaca, Tompkins County, X 0.6.

2-3. Manticoceras sinuosum tardum (Clarke, 1898) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 130NYSM 3804, ventral and lateral views of the lectotype, selected herein, a plastotype of the surviving syntype figuredby Clarke (1898: pl. 6, fig. 31) recorded as from the Naples Beds at Naples and by Miller (1938: 123) as from theCashaqua Shale in Briggs Gully (Loc. 43), Honeoye Lake, Ontario County, X 3.7.

4-5. Manticoceras simulator (Hall, 1874) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 124NYSM 3797, ventral and lateral views of the holotype figured by Hall (1876: pl. 69, figs 1-2) and recorded as fromIthaca, Tompkins County; probably from the Ithaca Formation of the Genesee Group, X 1.2.

6-8. Manticoceras sinuosum sinuosum (Hall, 1843) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 124NYSM 3762, enlarged lateral view and lateral and ventral views of a specimen figured by Clarke (1898: pl. 1, fig. 10,pl. 4, fig. 16) asMant. Pattersoni, recorded as from "a concretion in the soft shales at Naples, N.Y.," and thought tobe from the Cashaqua Shale. 6, X 1.6; 7, X 0.8; 8, X 0.6.


PLATE 10

Figure Page1-12. Manticoceras sinuosum sinuosum (Hall, 1843) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 124

1-2. NYSM 13805 (CUMP 42004), apertural and lateral views of specimen collected by Donald Zenger from theParrish Limestone, Cashaqua Shale, Conklin Gully (Loc. 47/2), near Naples, Ontario County, X 0.8.

3-4. NYSM 13817, apertural and lateral views of pyritic specimen from the Cashaqua Shale, Beards Creek (Loc.38/2), Pine Tavern, Livingston County, X 0.8.

5-7. NYSM 13827, lateral, apertural, and ventral views of baritic specimen from the Shurtleff Septarian Horizon,upper Cashaqua Shale, Shurtleff's Gully (Loc. 41/1), near Livonia, Livingston County, X 1.2.

8-9. NYSM 13835, apertural and lateral views of baritic specimen from the Shurtleff Septarian Horizon, upperCashaqua Shale, North McMillan Creek (Loc. 41a/5), Conesus Lake, Livingston County, X 0.8.

10. NYSM 13875, lateral view of specimen from Cashaqua Shale, Randall Gully (Loc. 44/4), near Bristol Center,Ontario County, X ca. 0.6.

11. NYSM 13823, lateral view of a baritic specimen from the Shurtleff Septarian Horizon, upper Cashaqua Shale,Shurtleff's Gully (Loc. 41/1), near Livonia, Livingston County, X 3.2.

12. NYSM 13806, lateral view of specimen from the Parrish Limestone, Cashaqua Shale, Rumpas Hill (Loc. 47a/1),near Naples, Yates County, X ca. 0.5.




PLATE 11

Figure Page1-13. Manticoceras sinuosum sinuosum (Hall, 1843) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 124

All baritized specimens from Shurtleff Septarian Horizon, upper Cashaqua Shale.1. NYSM 13876, lateral view of specimen from Shurtleff's Gully (Loc. 41/1), near Livonia, Livingston County, X ca.

4.6.2. NYSM 13877, lateral view of specimen from Shurtleff's Gully (Loc. 41/1), near Livonia, Livingston County, X

3.2.3, 7. NYSM 13878, lateral and ventral views of a specimen from Cottonwood Point (Loc. 40/13), Conesus Lake, Liv-

ingston County, X ca. 12.4-5. NYSM 13818, ventral and lateral views of specimen from Shurtleff's Gully (41/1), near Livonia, Livingston

County, X ca. 11.6. NYSM 3755, lateral view of hypotype figured by Clarke (1898: pl. 1, fig. 1) from the Portage (Naples) beds, Ho-

neoye Lake; probably from the Shurtleff Septarian Horizon, X ca. 13.8. NYSM 13819, lateral view of a specimen from Shurtleff's Gully (Loc. 41/1), near Livonia, Livingston County, X

ca. 9.6.9-10. NYSM 13828, lateral and apertural views of a specimen from Shurtleff's Gully (Loc. 41/1), near Livonia, Liv-

ingston County, X 1.2.11. NYSM 13825, lateral view of a specimen from from Shurtleff's Gully (Loc. 41/1), near Livonia, Livingston

County, X ca. 12.12. NYSM 3756, lateral view of specimen figured by Clarke (1898: pl. 1, fig. 2) from the Portage (Naples) beds, Ho-

neoye Lake; probably from the Shurtleff Septarian Horizon, X ca. 12.13. NYSM 3758, lateral view of a specimen figured by Clarke (1898: pl. 1, fig. 4) from the Portage (Naples) beds,

Honeoye Lake; probably from the Shurtleff Septarian Horizon, X ca. 15.


PLATE 12

Figure Page1-7, 11. Manticoceras sinuosum clausium n. ssp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 129

1-2. NYSM 13857, lateral and ventral views of recrystallized internal mold with septa and shell wall partially baritizedfrom the lower Cashaqua Shale, Whetstone Brook (Loc. 42/5), near Honeoye, Ontario County. 1, X 0.8; 2, X1.6.

3-4. NYSM 13839, lateral view of the holotype with the same preservation as specimen in Figs 1-2, from the lowerCashaqua Shale, Randall Gully (Loc. 44/3), near Bristol Center, Ontario County, X 1.2.

5. NYSM 13846, lateral view of specimen with the same preservation from the same horizon and locality (Loc. 44/3)as that in Figs 3-4, X 2.4.

6. NYSM 13858, lateral view of specimen with the same preservation as that in Figs 1-2 from the lower CashaquaShale, Whetstone Brook (Loc. 42/5), near Honeoye, Ontario County, X 1.6.

7, 11. NYSM 13865, lateral and ventral view of baritized specimen from the same horizon and locality (Loc. 42/5) asthat in Fig 6, X ca. 12.

8-10. Manticoceras sinuosum sinuosum (Hall, 1843) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 124NYSM 13829, lateral, ventral, and apertural views of baritized specimen from the Shurtleff Septarian Horizon, upperCashaqua Shale, Shurtleff's Gully (Loc. 41/1), near Livonia, Livingston County, X 1.2.




PLATE 13

Figure Page1-15. Manticoceras sinuosum clausium n. ssp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 129

All specimens are from lower Cashaqua Shale at Whetstone Brook (Loc. 42/5), near Honeoye, Ontario County, ex-cept those in Figs 7, 9, and 12 from Randall Gully (Loc. 44/3), near Bristol Center, Ontario County.1. NYSM 13863, lateral view, X ca. 11.6.2-3. NYSM 16559, lateral and ventral views. 2, X ca. 11; 3, X ca. 12.4. NYSM 16560, lateral view of protoconch and first whorl of a specimen showing constriction, X ca. 14.5-6. NYSM 13863, lateral and ventral views. 5, X ca. 11; 6, X ca. 12.7. NYSM 13841, lateral view, X 2.4.8. NYSM 13859, lateral view, X 2.9. NYSM 16562, lateral view, X ca. 6.4.10-11. NYSM 13866, lateral and ventral views, X ca. 12.12. NYSM 13846, lateral view, X 1.6.13. NYSM 13865, apertural view, X ca. 11.2.14-15. NYSM 13868, views showing the dorsal wrinkle, X ca. 11.2.


PLATE 14

Figure Page1-10. Manticoceras sinuosum tardum Clarke, 1898 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 130

All specimens from the lower Cashaqua Shale.1. NYSM 13879, enlarged view of early whorls based on a specimen from Briggs Gully (Loc. 43/3), Honeoye Lake,

Ontario County, X ca. 12.2, 4, 6-7. NYSM 13874, coarse ribbing of early whorls based on a specimen from Briggs Gully (Loc. 43/3), Honeoye

Lake, Ontario County. 2, X 3.2; 4 (oblique view), X ca. 15; 6, X ca. 7.7; 7 (oblique view), X ca. 9.3. NYSM 3766, lateral view of a specimen figured by Clarke (1898: pl. 2, fig. 3) asManticoceras pattersoni and show-

ing the protoconch; recorded as from the Naples Beds, Honeoye Lake (not Naples, New York) probably from theCashaqua Shale, Briggs Gully (Loc. 43), Ontario County, X 11.

5. NYSM 13880, growth lines of an older specimen from Briggs Gully (Loc. 43/3), Honeoye Lake, Ontario County,X 3.2.

8, 10. NYSM 16541, ribbing in early stages based on a specimen from Briggs Gully (Loc. 43/3). Honeoye Lake, On-tario County, X ca. 20.

9. NYSM 16561, growth lines of middle whorls, based on a specimen from Briggs Gully (Loc. 43/3), Honeoye Lake,Ontario County, X 3.2.




PLATE 15

Figure Page1-2. Manticoceras sinuosum sinuosum (Hall, 1843) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 124

MRH Collection, baritized replacement of the shell showing dorsal wrinkle layer and its confluence with the ventralwrinkle layer; based on a specimen collected by J. W. Wells from the Cashaqua Shale, Shurtleff Septarian Horizon,Shurtleff Gully (Loc 41), near Livonia, Livingstone County, X 5.


PLATE 16

Figure Page1-2, 9-10.Manticoceras lamed (G. & F. Sandberger, 1850) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 132

1-2. NYSM 12066, lateral and ventral views of a specimen from the Angola Shale, Lake Erie shore at the mouth ofFarnham Creek (Loc. 71/9a), Erie County, X 1.6.

9-10. NYSM 12057, ventral and lateral views of a specimen from equivalents of the Angola Shale in Wolf Creek (Loc.85), Castile, Wyoming County, X 0.8.

3-8. Manticoceras aff. lamed (G. & F. Sandberger, 1850) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 136All specimens from the Point Breeze Goniatite Bed, Angola Shale, in Relyea Creek (Loc. 82/6), Wyoming County.3-4. NYSM 12068, ventral and lateral views, X 1.6.5-6. NYSM 12070, lateral and ventral views, X 2.4.7-8. NYSM 12060, lateral and ventral views, X 2.4.

11-12. Manticoceras sinuosum sinuosum (Hall, 1843) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12411. MRH Collection, lateral view of a baritic replacement showing an extension of the flanks and umbilical shoulder

of one whorl onto the flanks of the preceding whorl; specimen collected by J. W. Wells from the Shurtleff Septar-ian Horizon, Cashaqua Shale, in Shurtleff Gully (Loc. 41), Livingston County: X 2.

12. NYSM 3761, lateral view of a baritic replacement figured by Clarke (1898: pl. 1, fig. 9) probably from the Shurtl-eff Septarian Horizon, Cashaqua Shale, at a locality near Honeoye Lake, Ontario County, X 5.4.




PLATE 17

Figure Page1. Archoceras wabashense (Kindle, 1901) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 85

GSC 5388, lateral view of a paratype from the lower New Albany Shale, about 0.75 mi (1.2 km) east of Delphi, In-diana, and just west of the crossing of the Monon Railroad and the Camden Pyke, X 3.2.

2-3. Clauseniceras delphiense (Kindle, 1901) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 852. GSC 5343a, lateral views of a paratype from same locality as specimen in Fig. 1, X 4.3. GSC 5343b, lateral views of a paratype from same locality as specimen in Fig. 1, X 3.2.

4. Manticoceras sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 137NYSM 3730, lateral view of a specimen figured by Clarke (1898: pl. 6, fig. 10) asManticoceras accelerans andrecorded as from the "soft shales at Naples," Ontario County, X 3.2.

5-10. Manticoceras sinuosum apprimatum Clarke (1898) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1235. NYSM 3731, lateral view of a cast of a cotype figured by Clarke (1898: pl. 6, fig. 27) recorded as from the Naples

Beds at Griswold, Genesee County, and probably from the Genundewa Limestone exposed in Murder Creek(Loc. 14a), Genesee County, X 3.4.

6-7. NYSM 3733, lateral and apertural views of a cotype preserved as a baritic replacement figured by Clarke (1898:pl. 6, fig. 29) from the Genundewa Limestone at Middlesex, Yates County, X 2.8.

8-10. NYSM 3732, apertural, lateral, and cross sectional views of the lectotype from the same locality and horizon asthat in Figs 6-7, X 2.5.


PLATE 18

Figure Page1-2, 5. Koenenites? fasciculatus (Clarke, 1898) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 120

1-2. NYSM 3738, apertural and lateral views of a baritic replacement figured by Clarke (1898: pl. 6, fig. 21), said tobe from the Genundewa Limestone, Canandaigua Lake,Yates County, X 3.2.

5. NYSM 3737, lateral view of a baritic replacement figured with the cotypes by Clarke (1898: pl. 6, fig. 18), said tobe from the Genundewa Limestone on Canandaigua Lake, Yates County, X 3.8.

3-4, 6-9. Carinoceras sororium (Clarke, 1898) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1373-4. NYSM 3800, ventral and lateral views of a gutta percha mold figured by Clarke (1898: pl. 4, fig. 4) prepared

from a specimen from Big Sister Creek, Angola, Erie County, and probably from the lower Angola Shale, X 4.6-7. NYSM 3801. oblique and lateral views of a gutta percha mold figured by Clarke (1898: pl. 4, fig. 5) prepared

from a specimen from Big Sister Creek, Angola, Erie County, and probably from the lower Angola Shale, X 3.8.8-9. NYSM 3802. lectotype here designated, ventral and lateral views of a gutta percha mold figured by Clarke (1898:

76, text-fig. 56) prepared from a specimen from Big Sister Creek, Angola, Erie County, and probably from thelower Angola Shale, X 3.8.

10. Delphiceras unduloconstrictum (Miller, 1938) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 141GSC 5346, lateral view of the holotype figured by Miller (1938: pl. 22, fig. 7) from the lower New Albany Shale,about 0.75 mi (1.2 km) east of Delphi, Indiana, and just west of the crossing of the Monon Railroad and the Cam-den Pyke, X 3.4.

11-12. Manticoceras nodifer (Clarke, 1885) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 123USNM 14994b, lateral view of the lectotype (designated by Becker & House, 1993: 132, pl. 2, figs 2-3) and an en-largement to show the nodosity of the inner whorls, based on a specimen figured by Clarke (1898: pl. 6, fig. 25)from the Genundewa Limestone, Canandaigua Lake, Yates County. 11, X 2.4; 12, X 10.




PLATE 19

Figure Page1-9. Delphiceras cataphractum (Clarke, 1898) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 141

1. NYSM 3643, lateral view of the lectotype here designated, a cotype figured by Clarke (1898: pl. 6, fig. 8) from theHanover Shale near Java, Wyoming County, and probably from nodules in the lower Hanover Shale in BeaverMeadow Creek (Loc. 92), X 4.

2-3. NYSM 3641. two views of a block showing several baritic shell replacements, one of which was figured by Clarke(1898: pl. 6, fig. 6) from the same locality and horizon as the specimen in Fig. 1, X 3.8.

4. NYSM 3644, lateral view of a cotype figured by Clarke (1898: pl. 6, fig. 9) from the same locality and horizon asthe specimen in Fig. 1, X 4.




8-9. NYSM 3638, ventral and lateral views of a cotype figured by Clarke (1898: pl. 6, fig. 3) from the same localityand horizon as the specimen in Fig. 1, X 4.

10-16. Manticoceras contractum Clarke, 1898 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12310-11. NYSM 3734, lateral and ventral views of a cotype figured by Clarke (1898: pl. 6, fig. 1) from the Genundewa

Limestone near Middlesex, Yates County, X 8.12-15. NYSM 3735, lateral and ventral views of the lectotype, here designated, a cotype figured by Clarke (1898: pl.

6, fig. 2) from the Genundewa Limestone, Canandaigua Lake. 12-13, X 2; 14-15, X 4.16. NYSM 5125, cross section of a probable cotype figured by Clarke (1898: text-fig. 44), said by Miller (1938: 82)

to be from the same locality and horizon as the lectotype in Figs 12-15, X 3.2.


PLATE 20

Figure Page1-12. Sphaeromanticoceras rhynchostomum (Clarke, 1898) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 142

1-2. NYSM 12078, oblique and lateral stereoscan photographs of a baritic shell replacement of inner whorls based ona specimen from the Point Breeze Goniatite Bed, Angola Shale, in Hampton Brook (Loc. 74/6b), Erie County. 1,X 8; 2, X 16.

3-6. NYSM 12079, oblique, ventral, and apertural views of a baritic shell replacement of inner whorls of a specimenfrom the same locality and horizon, X 16.

7-8. NYSM 12132 (6680 I), lateral and apertural views of a specimen collected by L. V. Rickard from upperRhinestreet Shale equivalents in Relyea Creek (Loc. 60), Wyoming County, X 1.6.

9-10. NYSM 12130, lateral and ventral views of a specimen from the Point Breeze Goniatite Bed, Angola Shale in theWest Branch of Cazenovia Creek (Loc. 75/6), Erie County, X 2.4.

11-12. NYSM 12131, ventral and lateral views of a specimen from the Angola Shale in Hampton Brook (Loc. 74/6a),Erie County, X 2.4.




PLATE 21


1-2. NYSM 3789, ventral and lateral views of a gutta percha mold figured by Clarke (1898: pl. 4, fig. 11) preparedfrom a specimen from Big Sister Creek (Loc. 73), Angola, Erie County, and probably from the lower AngolaShale, X 3.8.

3-4. NYSM 3786, lateral and ventral views of a gutta percha mold figured by Clarke (1898: pl. 4, fig. 8) preparedfrom a specimen from the same locality and horizon as that in Figs 1-2, X 4.

5. NYSM 3794, cross section of a specimen figured by Clarke (1898: text-fig. 41) based on a specimen from the samelocality and horizon as that in Figs 1-2, X 0.8.

6-8. NYSM 3790, ventral, lateral, and apertural views of a gutta percha mold figured by Clarke (1898: pl. 4, fig. 12)based on a specimen from the same locality and horizon as that in Figs 1-2, X 3.8.

9, 11. NYSM 3788, apertural and lateral views of a gutta percha mold figured by Clarke (1898: pl. 4, fig. 10) (speci-men on left of Fig. 11) based on a specimen from the same locality and horizon as that in Figs 1-2, X 3.8.

10. NYSM 3785, lateral view of a gutta percha mold figured by Clarke (1898: pl. 4, fig. 7) (specimen on left of Fig.10 only) based on a specimen from the same locality and horizon as that in Figs 1-2, X 4.

12. NYSM 3784 and 3787, lateral view (bottom left hand corner) of NYSM 3784 figured by Clarke (1898: pl. 4, fig.6) and oblique view (largest specimen close to NYSM 3784) of NYSM 3787, a specimen figured by Clarke(1898: pl. 4, fig. 9); the entire block is a gutta percha mold prepared from the same locality and horizon as that ofFigs 1-2, X 3.7.


PLATE 22


1-3. NYSM 12089, ventral, lateral, and cross sectional views of a specimen from the lower Angola Shale in HamptonBrook (Loc. 74/6a), Erie County, X 0.8.

4-5. NYSM 3793, lateral and ventral views of a specimen figured by Clarke (1898: text-fig. 30) labeled as from theNaples Beds at Angola, and probably from the lower Angola Shale at Big Sister Creek (Loc. 73), Angola, ErieCounty, X 0.8.

6-7. NYSM 3796, lectotype, here designated, a cotype figured by Clarke (1898: 68, text-fig. 43) probably from thelower Angola Shale and labeled as from Big Sister Creek (Loc. 73), Angola, Erie County, X 0.8.

8. NYSM 12035, lateral view of a specimen from the lower Angola Shale in Hampton Brook (Loc. 74/6a), ErieCounty, X 0.8.

9. NYSM 12085, cross section showing the form of the early whorls and an asymmetrically placed nepionic whorlbased on a specimen from upper Rhinestreet Shale equivalents in Johnson Creek (Loc. 58/1), Wyoming County,X 6.4.




PLATE 23

Figure Page1-4. Carinoceras aff. vagans (Clarke, 1898) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 141

1-2. NYSM 12044, lateral and ventral views of a specimen from the lower Angola Shale equivalents in Glade Creek(Loc. 77/1), Wyoming County, X 0.8.

3-4. NYSM 12043, ventral and lateral views of a specimen from the lower Angola Shale equivalents at Varysburg(Loc. 79/10), Wyoming County, X 0.8.

5-16. Carinoceras vagans (Clarke, 1898) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1385-6. NYSM 12048, ventral and lateral views of a specimen from the Relyea Creek Horizon, Rhinestreet Shale, Relyea

Creek (Loc. 60/3), Wyoming County, X 1.6.7-8. NYSM 12047, ventral and lateral views of a specimen from just below the Point Breeze Goniatite Bed, lower An-

gola Shale, in Hampton Brook (Loc. 74/6a), Erie County, X 1.6.9-10. NYSM 12041, ventral and lateral views of a specimen from the Point Breeze Goniatite Bed, Angola Shale, in

Cazenovia Creek (Loc. 75/6), Erie County, X 2.4.11-12. NYSM 12036, ventral and lateral views of a specimen from the Point Breeze Goniatite Bed, Angola Shale, in

Big Sister Creek (Loc. 73/6), Erie County, X 1.6.13-14. NYSM 3805, lateral and ventral views of holotype figured by Clarke (1898: pl. 6, figs 11-12) recorded as from

"a loose block of sandstone among Portage outcrops in the town of Naples" and said to be from the Grimes Silt-stone, a Rhinestreet Shale equivalent, X 1.6.

15-16. NYSM 12133, lateral and ventral views of a crushed specimen from lower Angola Shale equivalents in StonyCreek (Loc. 83/8), Warsaw, Wyoming County, X 1.2.


PLATE 24

Figure Page1-4. Acanthoclymenia genundewa (Clarke, 1898) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 105

1-2. NYSM 3646, syntype at 5.57 mm diameter, figured by Clarke (1898: pl. 8, fig. 2) from the Genundewa Lime-stone, probably at Genundewa Point (Loc. 23b), Canandaigua Lake, Yates County, X 8.

3. NYSM 3647, syntype at 6.3 mm diameter, from the Genundewa Limestone, possibly from Bristol, OntarioCounty, figured by Clarke (1898: pl. 8, fig. 3), possibly also from Genundewa Point, Canandaigua Lake, as statedon the caption but a label indicating Middlesex, X 6.

4. NYSM 3645, syntype, here designated lectotype, figured by Clarke (1898: pl. 8, fig. 1) from the GenundewaLimestone at Genundewa Point (Loc. 23b), Canandaigua Lake, Yates County, X 8.

5-9. Probeloceras lutheri (Clarke, 1885) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1565, 7. NYSM 4067, cast of the holotype figured by Clarke (1885: pl. 2, fig. 8; 1898: pl. 7, fig. 8), with a possible addi-

tional cast, X 2.6, 9. NYSM 4066 (formerly 12480/6), figured by Clarke (1898: pl. 7, fig 7), recorded as from Naples, and probably

from the shale (Cashaqua Shale) overlying the Parrish Limestone in Conklin (Parrish) Gully (Loc. 47/2), nearNaples, Ontario County. 6, X 2.4; 9, X 1.6.

8. USNM 14993, cast of the holotype figured by Clarke (1885: pl. 2, fig. 8). The USNM label records the original asfrom "shales which overlie the concretionary lst of Parrish Gully," that is, from the shale (Cashaqua Shale) overly-ing the Parrish Limestone in Conklin (Parrish) Gully (Loc. 47/2), near Naples, Ontario County, X 1.6.




PLATE 25

Figure Page1. Probeloceras lutheri (Clarke, 1885) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 156

NYSM 3648, lateral view of the holotype of Gephyroceras holzapfeli Clarke, 1898, figured by Clarke (1898: pl. 7, fig.17), a crushed pyritic specimen said to have come from the Cashaqua Shale at Eighteenmile Creek, Erie County, X3.6.

2-5. Naplesites iynx (Clarke, 1898) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 161All specimens probably from the lower Rhinestreet Shale around Naples, Yates County, although they have been la-beled as from the Cashaqua Shale.2. NYSM 3582, lateral view of a cotype figured by Clarke (1898: pl. 7, fig. 12), X 5.3. NYSM 3586, lateral view of a cotype figured by Clarke (1898: pl. 7, fig. 16), X 3.4. NYSM 3583, lateral view of a cotype showing the early whorls figured by Clarke (1898: pl. 7, fig. 13), X 5.5. NYSM 3581 (formerly 12060/1), specimen designated lectotype by Yatskov (1990); lateral view of a cotype figured

by Clarke (1898: pl. 7, fig. 11), X 4.4.

6. Naplesites naplesense (Clarke, 1898) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 160NYSM 4072, holotype of Probeloceras? naplesense (Clarke, 1898), sometimes assigned to Neomanticoceras. Designatedtype species of Naplesites by Yatskov (1990). Figured by Clarke (1898: pl. 7, fig. 18), and probably from theRhinestreet Shale at Naples, Yates County, X 1.2.


PLATE 26

Figure Page1-4, 6. Tornoceras uniangulare uniangulare (Conrad, 1842) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 163

1-2. AMNH 5476/1, lateral and apertural views of the holotype, the only specimen figured by Conrad (1842: pl. 16,fig. 4), from the Leicester Pyrite, near Leicester, Livingston County, X 2.

3-4. NYSM 13097 (CU 40115a), ventral and lateral views of a specimen from the Leicester Pyrite, near Leicester, Liv-ingston County, X 1.8.

6. NYSM 13098 (CU 40115b), lateral view of a specimen from the Leicester Pyrite, near Leicester, LivingstonCounty, X 2.4.

5. Tornoceras uniangulare widderi House, 1965 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 81USNM 137701, holotype from the Widder Shale, No. 4 Hill, Aux Sables River, ca. 2 mi (3.2 km) north of Arkona,Southern Ontario, collected by G. A. Cooper and P. E. Cloud, X 1.1.

7-10. Tornoceras arcuatum House, 1965 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1657. NYSM 12051, lateral view of a specimen collected by WTK from the Crosby Sandstone in Willow Grove Creek

(Loc. E, Py-14/1), East Branch of Keuka Lake, Schuyler County, X 1.6.8-9. NYSM 12675, apertural and lateral views of a topotype collected by MRH from the Squaw Bay Limestone at

Partridge Point, Alpena, Michigan, X 2.4.10. NYSM 12024, lateral view of a specimen collected by WTK from the Crosby Sandstone in Sunset Point Gully

(Loc. F, Py-16/1), East Branch of Keuka Lake, Yates County, X 1.6.




PLATE 27

Figure Page1-6. Tornoceras aff. uniangulare (Conrad, 1842) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 164

1-2. NYSM 12641 (MRH D1425), lateral and ventral views of a specimen collected by MRH from the upper Win-dom Shale at Grove's Creek Quarry (Loc. 4), southeast of Ovid, Seneca County, X 3.6.

3-4. NYSM 12643 (MRH D1441), lateral and ventral views of a specimen collected by MRH from the upper Win-dom Shale at Grove's Creek Quarry (Loc. 4), southeast of Ovid, Seneca County, X 3.2.

5-6. NYSM 12642 (MRH D1442), lateral and ventral views of a specimen collected by MRH from upper WindomShale at Grove's Creek Quarry (Loc. 4), southeast of Ovid, Seneca County, X 3.2.

7-10. Truyolsoceras bicostatum (Hall, 1843) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 179NYSM 11957 (MRH D1364), lateral and ventral views of the early whorls showing the nepionic constriction andcommencement of ventrolateral furrows of a specimen collected by MRH from the Corell's Point Goniatite Bed atCorell's Point (Loc. 97), near Brocton, Chautauqua County, X 16.

11. Tornoceras? sp. House, 1965 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11NYSM 11633 (5733), lateral view of a specimen from the West Brook Member of the Tully Limestone from nearBorodino (Loc. 6), Onondaga County, X 0.8.

12. Linguatornoceras aff. linguum (G. & F. Sandberger, 1851) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 167NYSM 12023, lateral view of a specimen from the upper Rhinestreet Shale equivalent in Johnson Creek (Loc. 58/1),Wyoming County, X 2.4.

13-14. Linguatornoceras linguum (G. & F. Sandberger, 1851) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16713. Schlüter Collection, Geologisch-Paläontologisches Institut, University of Bonn, West Germany; lateral view of a

specimen recorded as from the Upper Devonian, and probably UD I-G, at Büdesheim, Germany, X ca. 2.14. Schlüter Collection, Geologisch-Paläontologisches Institut, University of Bonn, Germany, lateral view of a speci-

men recorded as from the Upper Devonian, and probably UD I-G, at Büdesheim, Germany, X 2.

15. Tornoceras cf. typum (G. & F. Sandberger, 1851) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 166NYSM 4093, lateral view of a specimen from the “Naples formation” at Mount Morris, Livingston County, figuredby Hall (1879: pl. 72, figs 6-7), X 1.2.

16. Tornoceras arcuatum House, 1965 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 165NYSM 12671 (MRH D1440), lateral view of a topotype collected by MRH from the Squaw Bay Limestone nearPartridge Point, Alpena, Michigan, X 2.4.


PLATE 28

Figure Page1-5. Tornoceras uniangulare compressum Clarke, 1897 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 164

1-2. NYSM 12125, lateral views of a specimen collected by MRH from the Genundewa Limestone from the embank-ment locality at Bethany Center (Loc. 24), Genesee County. 1, X 1.2; 2, X 3.2.

3. NYSM 12126, lateral view of a specimen collected by MRH from the Genundewa Limestone from the embank-ment locality at Bethany Center (Loc. 24), Genesee County, X 1.6.

4-5. NYSM 12127, ventral and lateral views of a specimen collected by MRH, the ventral view showing the distinc-tive blunt venter of the subspecies; from the Genundewa Limestone from the embankment locality at BethanyCenter (Loc. 24), Genesee County, X 1.2.

6-10. Tornoceras uniangulare obesum Clarke, 1897 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1656-7, 10. NYSM 16564, ventral and lateral views of a baritized specimen from the Shurtleff Septarian Horizon of the

upper Cashaqua Shale in Shurtleff ’s Gully (Loc. 41) near Livonia, Livingston County. 6-7, X 0.8; 10, X 1.6.8-9. NYSM 11263 (E 546(ii)), ventral and lateral views of a baritized specimen showing the dorsal wrinkle layer;

from the Shurtleff Septarian Horizon, upper Cashaqua Shale, probably in the area of Honeoye Lake, OntarioCounty, X 3.9.




PLATE 29

Figure Page1-15. Phoenixites concentricus (House, 1965) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 173

1, 4-5. NYSM 11964 (6679E), ventral, lateral, and apertural views of a specimen from the Corell's Point GoniatiteBed at Corell's Point (Loc. 97), near Brocton, Chatauqua County, X 1.6.

2-3. NYSM 11963 (6679D), lateral and ventral views of the holotype from the Corell's Point Goniatite Bed,Gowanda Shale, at Corell's Point (Loc. 97), near Brocton, Chautauqua County, X 1.2.

6-9. NYSM 11965 (MRH D1401), lateral and ventral views of early whorls showing the nepionic constriction anddevelopment thereafter of ventrolateral furrows, based on a specimen from the Corell's Point Goniatite Bed atCorell's Point (Loc. 97), near Brocton, Chautauqua County, X 16.

10. NYSM 12549, enlargement of the dorsal lobe of a specimen from the Corell's Point Goniatite bed, GowandaShale, at Corell's Point (Loc. 97), near Brocton, Chatauqua County, X 3.2.

11. NYSM 16565, lateral view of a specimen thought to come from Gowanda Shale equivalents at Java (Loc. 109),Wyoming County, X 2.4.

12. NYSM 16566, lateral view of a specimen showing the dorsal wrinkle layer from the same locality and horizon asthat in Fig. 11, X 1.6.

13. NYSM 11243, lateral view of the largest specimen of the species known; from the Corell's Point Goniatite Bed atCorell's Point (Loc. 97), near Brocton, Chautauqua County, X 4.

14-15. NYSM 16591, ventral and lateral views of a specimen showing a displaced siphuncle from the same localityand horizon as that in Fig. 13, X 2.4.


PLATE 30

Figure Page1, 3. Truyolsoceras clarkei (Miller, 1938) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 180

1. NYSM 5652, lateral view of a cotype figured by Miller (1938: pl. 14, fig. 15, copied from Clarke, 1904: text-fig.15-left) from Forestville (Loc. 99) Chautauqua County, from the interval of Gowanda Shale, Laona Siltstone orWestfield Shale, X 0.8.

3. NYSM 5654, lateral view of a cotype figured by Miller (1938: pl. 14, fig. 16, copied from Clarke, 1904: text-fig.15-right) from Forestville (Loc. 99), Chautauqua County, from the interval of Gowanda Shale, Laona Siltstone orWestfield Shale, X 1.2.

2, 4-14. Truyolsoceras bicostatum (Hall, 1843) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1792. AMNH 5888/1, lateral view of the only cotype originally figured by Hall (1843: 245, text-fig. 107 (8)), here desig-

nated lectotype, said to be from the shore of Lake Erie and thought to be from the Corell's Point Goniatite Bed atCorell's Point (Loc. 97), near Brocton, Chautauqua County, X 1.6.

4-6, 14. NYSM 11961, ventral and lateral views of a specimen collected by MRH from the Corell’s Point GoniatiteBed (Loc. 97), near Brocton, Chautauqua County. 4-6, X 2.4; 14, X 2.4.

7-8. NYSM 5659, ventral and lateral views of a specimen from the Corell's Point Goniatite Bed (Loc. 97), near Broc-ton, Chautauqua County, X 2.4.

9-11. NYSM 11958, apertural, lateral, and ventral views of a specimen from the same locality and horizon, X 2.4.12. NYSM 11238, lateral views of a specimen from the Corell's Point Goniatite Bed (Loc. 97), near Brocton, Chau-

tauqua County, X 1.2.13. AMNH 5888/1:1, lateral view of a specimen figured by Hall (1879: pl. 72, fig. 8, pl. 74, fig. 1) thought to be

from the Corell's Point Goniatite Bed (Loc. 97), near Brocton, Chautauqua County, X 1.6.

15. Truyolsoceras cf. bicostatum (Hall, 1843) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 180NYSM 12120, lateral view of a specimen from the Gowanda Shale from near the Mixer Road bridge over WalnutCreek (Loc. 99a) at Forestville, Chautauqua County, X 1.6.




PLATE 31

Figure Page1. Truyolsoceras cf. bicostatum (Hall, 1843) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 180

NYSM 12119, lateral view of a specimen from the Gowanda Shale from near the Mixer Road bridge over WalnutCreek (Loc. 99a) near Forestville, Chautauqua County, X 1.6.

2-3. Aulatornoceras auris (Quenstedt, 1846) Group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1762. NYSM 12114, oblique view of a specimen from the Angola Shale in Sheldon Creek (Loc. 78/4b), Erie County, X

1.6.3. NYSM 12117, fragment of a specimen from the lower Hanover Shale above Angel Falls, Java (Loc. 92/3),

Wyoming County, X 1.6.

4-6. Aulatornoceras aff. eifliense (Steininger, 1849) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1774. NYSM 12116, specimen showing distinctive bifid growth lines from the upper Rhinestreet Shale equivalents in

Kennedy Gulf (Loc. 63/1), Wyoming County, X 1.6.5-6. NYSM 12103, ventral and lateral views of a specimen from the Angola Shale on the shore of Lake Erie (Loc.

71/9a), at Point Breeze, Erie County, X 3.2.

7-13, 16-20. Aulatornoceras eifliense (Steininger, 1849) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1767. NYSM 12109, stereoscan lateral view of a specimen collected by WTK from the Cashaqua Shale in Buck Run

Creek (Loc. 39/17), Livingston County, X 8.8-9. NYSM 12108, ventral and lateral views of a specimen collected by WTK from the Cashaqua Shale in a tributary

of Cazenovia Creek (Loc. 32/9), Erie County, X 3.2.10-11. NYSM 12105, stereoscan oblique and lateral views of a specimen collected by WTK from the Cashaqua Shale

in the South Branch of Smoke Creek (Loc. 31/5), Erie County, X 8.12-13. NYSM 12110, stereoscan ventral and lateral views of a specimen collected by WTK from the Cashaqua Shale

in Murder Creek (Loc. 34/6), Genesee County, X 8.16-17. NYSM 12111, stereoscan lateral and ventral views of a specimen from the Cashaqua Shale in the South

Branch of Smoke Creek (Loc. 31/5), Erie County, X 8.18-20. NYSM 12107, lateral and ventral views of a baritized specimen collected by WTK from the Cashaqua Shale

(Shurtleff Septarian Horizon) in Shurtleff's Gully (Loc. 41/1), near Livonia, Livingston County, X 2.4.

14-15. Pharciceras? sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9214. NYSM 12128, lateral view of a specimen from the Geneseo Shale at Pleasant Valley, near Sherburne (Loc. 28),

Chenango County, X 1.2.15. NYSM 12129, lateral view of a specimen from the Geneseo Shale near Chaseville, Otsego County, X 1.2.


PLATE 32

Figure Page1. Aulatornoceras rhysum (Clarke, 1898) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 179

NYSM 4092, lateral view of the lectotype figured by Clarke (1898: text-fig. 100, pl. 8, fig. 14) labeled “HanoverShale, Java” and probably from the lower Hanover Shale in Beaver Meadow Creek at Java (Loc. 92), WyomingCounty, X 4.

2-13. Aulatornoceras paucistriatum (d'Archiac & de Verneuil, 1842) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1782-3. BMNS E 22464, ventral and lateral views of a specimen collected by I. G. Reimann probably from the lower An-

gola Shale, Hampton Brook (see Loc. 74), Erie County, X 3.2.4, 11. NYSM 12097, stereoscan lateral and edge views of a specimen from near the Point Breeze Goniatite Bed of the

Angola Shale in Hampton Brook (Loc. 74/6a), Erie County, X 8.5. NYSM 12098, stereoscan lateral view of a specimen from near the Point Breeze Goniatite Bed of the Angola Shale

in Hampton Brook (Loc. 74/6a), Erie County, X 8.6. NYSM 12099, stereoscan view of specimens on a block from near the Point Breeze Goniatite Bed of the Angola

Shale in Hampton Brook (Loc. 74/6a), Erie County, X 8.7-9. NYSM 12096, stereoscan ventral and lateral views of a specimen from the horizon (Loc. 74/6a) immediately

below the Point Breeze Goniatite Bed in Hampton Brook, Erie County, X 8.10. NYSM 12100, oblique view of a specimen from the Angola Shale of Hampton Brook (Loc. 74/3), Erie County, X

3.2.12-13. NYSM 12095, lateral and ventral views of a specimen from near the Point Breeze Goniatite Bed of the Angola

Shale in Hampton Brook (Loc. 74/6a), X 3.2.

14. Aulatornoceras aff. eifliense (Steininger, 1849) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 177NYSM 12102, lateral view of a specimen from near the Point Breeze Goniatite Bed of the Angola Shale in HamptonBrook (Loc. 74/6a), X 3.2.




PLATE 33

Figure Page1-2, 5-8. Linguatornoceras aff. linguum (G. & F. Sandberger, 1851) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 167

1-2. NYSM 11248, lateral and ventral views of a specimen collected by D. D. Luther in 1897 from Relyea Creek(Gibson's Glen) (Loc. 60, 82), Wyoming County, from the Rhinestreet or Angola Shale equivalents, X 2.1.

5-6. NYSM 12019, ventral and lateral views of a specimen from upper Rhinestreet Shale equivalents in TonowandaCreek at Varysburg (Loc. 57/2), Wyoming County, X 3.2.

7-8. NYSM 12020, ventral and lateral views of a specimen from the upper Rhinestreet Shale equivalents in JohnsonCreek (Loc. 58/1), Wyoming County, X 2.

3-4. Linguatornoceras sp., juvenile specimen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 167NYSM 12021, stereoscan lateral and ventral views of a specimen from the Point Breeze Goniatite Bed, Angola Shale,in the West Branch of Cazenovia Creek at Griffins Mills (Loc. 75/6), Erie County, X 8.

9-20. Crassotornoceras aff. crassum (Matern, 1931) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1689-10. NYSM 12013, stereoscan lateral and oblique views of a specimen from the Point Breeze Goniatite Bed, Angola

Shale, in the West Branch of Cazenovia Creek at Griffins Mills (Loc. 75/6), X 4.8.11-12. NYSM 12012, stereoscan ventral and lateral views of a specimen from the West Branch of Cazenovia Creek at

Griffins Mills (Loc. 75/6) from the level of the Point Breeze Goniatite Bed of the Angola Shale, X 8.13-14. NYSM 12010, stereoscan oblique and lateral views of a specimen from about the Point Breeze Goniatite Bed

of the Angola Shale in Hampton Brook (Loc. 74/6a), Erie County, X 8.15-16. NYSM 12015, stereoscan ventral and lateral views of a specimen from Angola Shale equivalents in Relyea

Creek, south of Warsaw (Loc. 82/6), Wyoming County, X 4.8.17-18. NYSM 12014, stereoscan ventral and lateral views of a specimen from Angola Shale equivalents in Johnson

Creek, near Earls (Loc. 81/6), Wyoming County, X 4.8.19-20. NYSM 12011, lateral and ventral views of a specimen from about the Point Breeze Goniatite Bed of the An-

gola Shale in Hampton Brook (Loc. 74/6a) south of Hamburg, Erie County, X 2.4.


PLATE 34

Figure Page1-4. Lobotornoceras aff. hassoni House, 1978 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 172

1. NYSM 12029, stereoscan lateral view of a specimen from the upper West River Shale in Seneca Point Creek (Loc.26a/3), Canandaigua Lake, Ontario County, X 4.8.

2-4. NYSM 12030, stereoscan ventral and oblique views of a specimen from the upper West River Shale in Whet-stone Brook (Loc. 25a/1), Ontario County, X 4.8.

5. Epitornoceras aff. peracutum (Hall, 1876) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 171CU 39652, lateral view of a specimen figured by Harris (1898: pl. 6, fig. 35) from the “middle Portage, near Ithaca,”Tompkins County, X 0.8.

6-7. Epitornoceras cf. peracutum (Hall, 1876) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 171NYSM/CU 369, lateral and ventral views of a specimen from the upper Geneseo Shale or lower Penn Yan Shale, FirTree Point, western side of Seneca Lake (Loc. 25), Yates County, X 0.8.

8. Epitornoceras peracutum (Hall, 1876) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 170NYSM 4091, lateral view of the holotype figured by Hall (1876: pl. 69, fig. 8, pl. 74, fig. 13; 1879: pl. 69, fig. 8, pl.74, fig. 13), probably from the Ithaca Formation at Ithaca, Tompkins County, X 0.8.




PLATE 35

Figure Page1-10. Cheiloceras (Cheiloceras) amblylobum (G & F. Sandberger, 1850) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 181

All specimens from the Corell's Point Goniatite Bed, lower Gowanda Shale, at Corell's Point (Loc. 97), Lake ErieShore, Chautauqua County.1-2. USNM 137666, lateral and apertural views of a specimen collected by W. Moran, X 1.6.3-5. NYSM/CU 40082, lateral and ventral views, X 1.6.6. MRH 251, lateral view, X 1.6.7-8. USNM 137665, ventral and lateral views of a specimen collected by W. Moran, X 1.6.9-10. NYSM/CU 40083, lateral and ventral views, X 1.6.

11, 13. Maeneceras aff. acutolaterale (G. & F. Sandberger, 1951) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 183USNM 137645, lateral view of a specimen collected by H. S. Williams in 1884 from the Ellicott Shale at PorterCreek, Summerdale (Loc. 110), Chautauqua County. 11, X 0.4; 12, X 0.7.

12. ?Genus edwinhalli (Clarke, 1898) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 183NYSM 4090, lateral view of the holotype figured by Clarke (1898: 111, text-fig. 85) from the Conneaut Formationat Nile, Allegany County, X 1.6.



Acanthoclymeniidae Schindewolf, 1955 . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . 72-73, 77, 98, 103, 114, 155; Text-figs 20-21

AcanthoclymeniaHyatt, 1900. . . . . . . . . . . . . . . . . 17, 21, 23, 29-30, 32,72-73, 82, 98, 102-104, 107, 109, 120, 157; Text-figs 21, 30A. forcipifer (G. & F. Sandberger, 1850) . . . . . . . . . . . . . 102, 247A. genundewa (Clarke, 1898) . . . . . . . . . . 79, 105-107, 155,

250; Text-figs 23, 30A-C, K, L; Table 6; Pl. 24, Figs 1-4A. aff. genundewa (Clark, 1898) . . . . . . . . . . . . . . . . . . . . . . 106A. cf. genundewa (Clarke, 1898) . . . . . . . . . . . . . . 105-106, 107A. neapolitana Clarke, 1892) . . . . . . . . . 35, 39-40, 79, 83, 103-

106, 214; Text-figs 23, 30E-H; Table 5; Pl. 6, Figs 1-15A. aff. neapolitana (Clarke, 1892) . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . 31, 33, 79, 83, 102-105; Text-figs 23, 30DA. cf. neapolitana (Clarke, 1892) . . . . . . . . . . . . . . . . . . . . 106A. orientale (Bogoslovsky, 1969) . . . . . . . . . . . . . . . . . . . . 102A. planorbe (G. & F. Sandberger, 1850) . . . . . . . . . . . . . . . 102A. sp. . . . . . . . . . . . . . . . . . . . . . . . . 79, 103, 107; Text-fig. 23A. sp. A Kirchgasser, 1975 . . . . . . . . . . . . . . . . . . . . . 31-32, 105

accelerans,Manticoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1, 137, 237acutiforme, Carinoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . 137-138, 140acutolaterale,Maeneceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86, 183, 185acutolaterale,Maeneceras aff. . . . . . . . . . . . . . . . 62, 79, 86, 182-183, 184-

185, 273; Text-figs 23, 61C-D, 62-63; Pl. 35, Figs 11, 13acutolateralis, Goniatites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 182acutum, Carinoceras . . . . . . . . . . . . . . . . . . . 137-138, 140; Text-fig. 44Bacutum, Oxytornoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 169acutum, Tornoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 169aequabile, Ponticeras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 98affine, Sphaeromanticoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 142, 157affinis, Goniatites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 142Afromaenioceras Göddertz, 1987 . . . . . . . . . . . . . . . . 73, 76; Text-fig. 21altaicum,Manticoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 150alveolatus, Prochorites . . . . . . . . . . . 35, 38-41, 79, 83, 109-110, 156-157,

185, 217; Text-figs 23, 31, 32C, 63; Table 7; Pl. 7, Figs 4-6alveolatus, Prochorites aff. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43, 49, 110amblylobum, Cheiloceras (Cheiloceras). . . . . . 59-62, 79, 86, 181-182, 185,

273; Text-figs 23, 59C, 61A-B; Table 35; Pl. 35, Figs 1-10Ammonites aequabile Beyrich, 1837 . . . . . . . . . . . . . . . . . . . . . . . . . . . 98Ammonites hoeninghausia von Buch, 1832 . . . . . . . . . . . . . . . . . . 112, 122Ammonites orbiculus Beyrich, 1837 . . . . . . . . . . . . . . . . . . . . . . . . . . . 142Ammonites primordalis von Buch, 1832 . . . . . . . . . . . . . . . . . . . . . . . . 142Ammonites simplex von Buch, 1832 . . . . . . . . . . . . . . . . . . . . . . . . . . . 163amplexum, Pharciceras . . . . . . . . . . . . . . . . 10-11, 79, 82, 89-90, 91, 185-

186, 205; Text-figs 23, 26D-H, 63; Table 2; Pl. 1, Figs 1-8Anarcestidae . . . . . . . . . . . . . . . 71-72, 73, 76, 98, 162; Text-figs 20-21anguisellatum, Naplesites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 160-161annissi, Crassatornoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 168apprimatum,Manticoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . 83, 123, 132apprimatum, Manticoceras sinuosum . . . . . . . . . . . 1, 29, 79, 83, 123, 132,

135, 185, 237; Text-figs 23, 35B, 42F, 63; Pl. 17, Figs 5-10archiaci, Hoeninghausia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 122archiaci, Hoeninghausia aff. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 83archiaci, Hoeninghausia cf. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 121Archoceras Schindewolf, 1937. . . . . . . . 79, 83, 85, 102, 149; Text-fig. 48

A. varicosum (Drevermann, 1901) . . . . . . . . . . . . . . . . . . . . . 85A. wabashense (Kindle, 1901) . . . . . . . . . . . . 237; Pl. 17, Fig. 1A. (Atlantoceras) sp. Kirchgasser & House, 1981. . . . . . . . . 102?A. sp. . . . . . . . 58-59, 79, 86, 155, 192; Text-figs 23, 48G, 63

Atlantoceras Bensaïd, 1974 . . . . . . . . . . . . . . . . . . . . . . . . . . . 76, 98, 102

arcuatum, Tornoceras . . . . . . . . . . 22, 79, 82, 158, 165-166, 17-, 254, 257;Text-figs 23, 54H-J, L; Pl. 26, Figs 7-10, Pl. 27, Fig. 16

arcuatum, Tornoceras cf. . . . . . . . . . . . . . 10, 158, 166, 170; Text-fig. 54Karkonense, Tornoceras uniangulare (House, 1965) . . . . . . . . . . . . . . . . . 81astarte, Goniatites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 163Atlantoceras Bensaïd, 1974 . . . . . . . . . . . . . . . . . . . . . . . . . . . 76, 93, 102Aulatornoceratini Becker, 1993. . . . . . . . . . . . . . . . . . . . . . . . . . . . 175Aulatornoceras Schindewolf, 1922 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . 39-40, 73, 86, 168, 175, 178-179; Text-fig. 21A. eifliense (Steininger, 1849) . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . 35, 38-40, 79, 83, 175-176, 177-178,265; Text-fig. 23; Table 31; Pl. 31, Figs 7-13, 16-20

A. aff. eifliense (Steininger, 1849) . . . . 49, 54, 79, 177, 265-266;Text-fig. 23; Table 32; Pl. 31, Figs 4-6, Pl. 32, Fig. 14

A. eifliense eifliense (Steininger, 1849) . . . . . . . . . . . . . . . . . . 177A. eifliense posterior Becker, 1993 . . . . . . . . . . . . . . . . . . . . 175A. auriforme (Oppenheimer, 1916) . . . . . . . . . . . . . . . . . . . . 176A. auris (Quenstedt, 1846) . . . . . . . . . . . . . . . . . . . . 55, 79, 85,

175-176, 178, 265; Text-figs 23, 60G; Pl. 31, Figs 2-3A. cf. auris (Quenstedt, 1846) . . . . . . . . . . . . . . . . . . . . . . . . 176A. auris bickensis (Wedekind, 1918) . . . . . . . . . . . . . . . . . . . 175A. constrictum (Steininger, 1853) . . . . . . . . . . . . . . . . . . . . . . 176A. keyserlingi (Müller, 1956) . . . . . . . . . . . . . . . . . . . . . . . . . 176A. lepiferum Becker, 1993 . . . . . . . . . . . . . . . . . . . . . . . . . . 175A. loeschmanni (Frech, 1902) . . . . . . . . . . . . . . . . . . . . . 169, 176A. paucistriatum (d’Archiac & de Verneuil, 1841) . . . . . . . . . . .

. . . . . . . . . . . . . . . . 54-55, 79, 85, 175-176, 178-179, 180,266; Text-figs 23, 60D, F; Table 33; Pl. 32, Figs 2-13

A. rhysum (Clarke, 1898) . . . . . . . . . . . . . . . . . . . 58-59, 79, 85,175-176, 178-179, 266; Text-figs 23, 60E; Pl. 32, Fig. 1

A. sandbergeri (Ford & Crick, 1897) . . . . . . . . . . . . . . . . . . . 175A. serriense Becker, 1993 . . . . . . . . . . . . . . . . . . . . . . . . . . . . 176

auriforme, Aulatornoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 176auris, Aulatornoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55, 79,

85, 175-176, 178, 265; Text-figs 23, 60G; Pl. 31, Figs 2-3auris, Aulatornoceras cf. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 176auris, Goniatites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 175-176auritum, Ponticeras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 98ausavense, Lobotornoceras . . . . . . . . . . . . 168, 171-172; Text-figs 58C, Fausavensis, Goniatites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 168barnetti, Pharciceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 186barroisi, Ponticeras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 98beckeri n. sp., Koenenites . . . . 1, 31-34, 79, 112-115, 117, 121-122, 185,

218; Text-figs 23, 33A-E, 34H, K, 63; Pl. 8, Figs 10-11, 14beckeri, n. sp., Koenenites, form D . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . 31, 33, 117, 122; Text-figs 33E, 34Kbelgicum, Tornoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 168Beloceratidae Hyatt, 1884 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . 72-73, 76-77, 94, 96, 98, 102-103, 155; Text-figs 20-21Beloceras Hyatt, 1884 . . . . . . . . 50, 73, 83-84, 94, 96, 155; Text-fig. 21

B. sagittarium (G. & F. Sandberger, 1851) . . . . 94; Text-fig. 27CB. williamsiWells, 1956 . . . . . . . . . . . . . . . . . . . . . . . . . . . 96

bicaniculatum, Lobotornoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 173bickensis, Aulatornoceras auris . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 175bicostatum, Tornoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 178bicostatum, Tornoceras (Aulatornoceras) . . . . . . . . . . . . . . . . . . . . . . . . 178bicostatum, Truyolsoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . 1, 59-61, 79, 86, 175-176, 178-179, 180, 257, 262; Text-figs 23, 59B, 60A-C, H; Table 34; Pl. 30, Figs 2, 4-13, 15

SYSTEMATIC INDEX

Pages denoting the start of principal discussions, locations of plate and text-figure illustrations, and tables of measurements are in bold font.


bicostatum, Truyolsoceras cf. . . . . . . . . . . . . . . . . . . . . . . . . . . . 59-61, 79,86, 180, 262, 265; Text-fig. 23; Pl. 30, Fig. 15, Pl. 31, Fig. 1

bidentatum, Pharciceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 89bilobatum, Falcitornoceras aff. . . . . . . . . . . . . . . . . . 172; Text-fig. 58Dbisulcatum, Ponticeras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 98buchii,Manticoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 129Carinoceras Lyashenko, 1957 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . 73, 77, 84, 122, 129, 137, 140; Text-figs 21, 42-44C. acutiforme (H. & G.Termier, 1950) . . . . . . . . . . 137-138, 140C. acutum (G. & F. Sandberger, 1850) . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . 137-138, 140; Text-fig. 44BC. galeatum (Wedekind, 1913) . . . . . . . . . . . . . . . . . . . 85, 140C. menneri Ljaschenko, 1957 . . . . . . . . . . . . . . . . . . . 137-138C. oxy (Clarke, 1897) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 144C. sororium (Clarke, 1898) . . . . . . . . . . . . . . . . . . . . . 1, 51, 79,

85, 137, 238; Text-figs 23, 42G; Pl. 18, Figs 3-4, 6-9C. vagans (Clarke, 1898) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . 43, 48, 54-55, 79, 84-85, 138-139, 140, 249; Text-figs 23, 43A-E, 44D, 46A; Table 16; Pl. 23, Figs 5-16

C. aff. vagans (Clarke, 1898) . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . 55, 79, 138, 141, 249; Text-fig. 23; Pl. 23, Figs 1-4

cataphractum,Delphiceras . . . . . . . . . . . . . . 1, 58-59, 79, 85, 134,141, 185, 241; Text-figs 23, 41G, 63; Pl. 19, Figs 1-9

cataphractum, Gephyroceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 141CeratobelocerasHouse & Kirchgasser, 1985 . . . . . . . . . . . . . . . 73, 77, 155Chaoceras Yatskov, 1991 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 161Cheiloceras Frech, 1897 . . . . . . 3, 59, 62, 73, 174, 181-182; Text-fig. 21

C. (Cheiloceras) amblylobum (G. & F. Sandberger, 1851) . . . . . .. . . . . . . . . . . . . . . . . . 59-62, 79, 174, 181-182, 185; Text-figs 23, 59C, 61A-B, 63; Table 35; Pl. 35, Figs 1-10

Cheiloceratidae Frech, 1897 . . . . . . . . . . . 72-73, 181; Text-figs 20-21Cheiloceratos Strand, 1929 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 181Cheilocerotes Strand, 1929 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 181Chemungense, Sandbergeroceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 97chemungense, Schindewolfoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50,

79, 84, 94, 96-97, 98, 206; Text-figs 23, 27I, 63; Pl. 2, Fig. 8chemungense, Schindewolfoceras aff. . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . 94, 97, 210; Text-fig. 27J; Pl. 4, Figs 1-4chemungensis, Goniatites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93Chiloceras Dreverman, 1901 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 181Chutoceras Becker & House 2000 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . 73, 98, 100, 107-109, 129; Text-fig. 21C. manticoides Becker & House, 2000 . . . . . . . . . . . . . . 107-109C. nundaium (Hall, 1874) . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . 1, 17, 23, 25-26, 77, 79, 82, 100, 107-108,109, 185, 221; Text-figs 23, 28H-K, 63; Pl. 9, Fig. 1

cinctum, Goniatites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 176clarkei, Tornoceras (Aulatornoceras) . . . . . . . . . . . . . . . . . . . . . . . . . . 61clarkei, Truyolsoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59,

61, 79, 86, 176, 179-180, 262; Text-fig. 23; Pl. 30, Figs 1, 3Clauseniceras Becker & House, 1993 . . . . . . . . . . . 73, 122; Text-fig. 21

C. delphiense (Kindle, 1901) . . . . . . . 85, 237; Pl. 17, Figs 2-3C. expectatum (Wedekind, 1913) . . . . . . . . . . . . . . . . . . . . . . 85

clausium n. ssp.,Manticoceras sinuosum . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . 1, 35, 40-41, 83, 129-130, 131, 226, 229; Text-figs 39A-I, 40A-B; Table 12; Pl. 12, Figs 1-7, 11, Pl. 13, Figs 1-15

clausum, Linguaornoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 166clausum, Tornoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 166ClymeniaMünster in Goldfuss, 1832 . . . . . . . . . . . . . 102-103, 107-108complanatus perlatus, Goniatites . . . . . . . . . . . . . . . . . . . . . . . . . . . 99, 108compressum, Tornoceras uniangulare . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . 27, 29, 79, 82, 157-158, 162, 164-165, 258;Text-figs 23, 53A-B, 54F, 55B; Table 24; Pl. 28, Figs 1-5

concentricum, Tornoceras (Tornoceras) . . . . . . . . . . . . . . . . . . . . . . . . 173concentricus, Phoenixites . . . . . . . . . . . . . . . . . . . . . . . . . 59-60, 86, 158,

173, 261; Text-figs 54M-O, 59A; Table 26; Pl. 29, Figs 1-15constrictum, Aulatornoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 176, 179contractum, Manticoceras . . . . . . . . . . . . . . . . . 1, 27, 29, 79, 83, 118, 123,

130, 132, 136-137, 241; Text-figs 23, 35B; Pl. 19, Figs 10-16cooperi, Koenenites . . . . . . . . . . . . . . . . . . . 112-113, 117, 120; Table 9cordatum,Manticoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . 129, 134, 136cordatum,Manticoceras lamed aff. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . 31, 83, 115, 136; Text-figs 33K-Lcordatum, Manticoceras lamed cf. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 83cordiforme, Crickites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 155cordiforme,Manticoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 129, 150Costamanticoceras Becker & House, 1993 . . . . . 73, 77, 122; Text-fig. 21

C. koeneni (Holzapfel, 1882) . . . . . . . . . . . . . . . . . . . . . . . 137costatum, Triainoceras . . . . . . . . . . . . . . . . . . . . . . . . 94; Text-fig. 27Ecostatus, Goniatites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 92CrassotornocerasHouse & Price, 1985 . . . 73, 157, 162, 168; Text-fig. 21

C. annissi House & Price, 1985 . . . . . . . . . . . . . . . . . . . . . 168C. ausavense crassum (Matern, 1931) . . . . . . . . . . . . . . . . . 168C. belgicum (Matern, 1931) . . . . . . . . . . . . . . . . . . . . . 168C. crassum (Matern, 1931) . . . . . . . . . . . . . . . . . . . . . . . 169C. aff. crassum (Matern, 1931) . . . . . . . . . . . . . 54-57, 85, 157,

168, 269; Text-figs 53E-F; Table 27; Pl. 33, Figs 9-20?C. isolatum Becker, 1993. . . . . . . . . . . . . . . . . . . . . . . . . . 168C. nitidum Becker, 1993 . . . . . . . . . . . . . . . . . . . . . . . . . . 168

crassum, Crassotornoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 169crassum, Crassotornoceras aff. . . . . . . . . . . . . . . . . . . . . 54-57, 79, 85, 157,

168, 269; Text-figs 23, 53E-F; Table 27; Pl. 33, Figs 9-20crassum, Crassoctornoceras ausavense . . . . . . . . . . . . . . . . . . . . . . . . . 168crassum, Tornoceras ausavense . . . . . . . . . . . . . . . . . . . . . . . . . . . 168Crickitinae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 122, 150CrickitesWedekind, 1913 . . . . . 73, 77, 122, 149-150, 155; Text-fig. 21

C. cordiforme (Miller, 1938) . . . . . . . . . . . . . . . . . . . . . . . . 155C. holzapfeli Wedekind, 1913 . . . . 150, 155; Text-figs 48D-FC. lindneri (Glenister, 1958) . . . . . . . . . . . . . . . . . . . . . . . . 58,

79, 85, 149-150, 151, 185; Text-figs 49, 63; Table 21?C. sp. juv. . . . . . 79, 86, 149, 155, 185; Text-figs 23, 48H, 63

Delphiceras Becker & House, 2000 . . 73, 77, 122, 134, 141; Text-fig. 21D. cataphractum (Clarke, 1898) . . . . . . . . 1, 58-59, 79, 85, 134,

141, 185, 241; Text-figs 23, 41G, 63; Pl. 19, Figs 1-9D. unduloconstrictum (Miller, 1938) . . 141, 238; Pl. 18, Fig. 10

delphiense, Clauseniceras . . . . . . . . . . . . . . . . . 85, 237; Pl. 17, Figs 2-3Delphinites Becker & House 1993 . . . . . . . . . . . . . . . . . . . . . . 122, 141Delphinites Sayn, 1901 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 141Devonopronorites . . . . . . . . . . . . . . . . . . . . . . . . . . 71, 72-73, 77, 89Devonopronoritidae Bogoslovsky, 1958 . . . . 72-73, 89; Text-figs 20-21discoidale, Ponticeras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 98domanicense, Ponticeras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 98Domanikoceras Becker & House, 1993 . . . . . . . . . . 73, 162; Text-fig. 21Edwin-halli, Tornoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . 62, 87, 183edwinhalli, ?Genus . . . . . . . . . . . . . . . . . . . . . . 183, 273; Pl. 35, Fig. 12Eidoprobeloceras Kirchgasser, 1968 . . . . . . . . . . . . . . . . . . . . . . . . . . 98

E. strix Kirchgasser, 1968 . . . . . . . . . . . . . . . . . . . . . . . . . . 109eifliense, Aulatornoceras . . . . . . . . . . . . . . . . . . . . . . . . . 35, 38-40, 79, 83,

175-176, 177-178, 265; Table 31; Pl. 31, Figs 7-13, 16-20eifliense, Aulatornoceras aff. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49, 54,

79, 177, 265-266; Table 32; Pl. 31, Figs 4-6, Pl. 32, Fig. 14eifliense, Tornoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 177eifliense eifliense, Aulatornoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . 177eifliense posterior, Aulatornoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . 175enfieldense n. sp. , Sandbergeroceras? . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . 1, 49-51, 93, 95, 206, 209; Pl. 2, Figs 1-2, Pl. 3, Figs 2-3


Enseites Becker & House, 1993 . . . . . . . . . . . . 73, 77, 98; Text-fig. 21E. sulcatum (Matern, 1931) . . . . . . . . . . . . . . . . . . . . . . . . 105

Eobeloceratidae . . . . . . . . . 71, 72-73, 76, 88-89, 155; Text-figs 20-21Epitornoceras Frech, 1902 . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16, 19-20,

73, 76-77, 82, 158, 162, 164, 166, 169-170, 171; Text-fig. 21E. mithracoides (Frech, 1887) . . . . 82, 169-170; Text-fig. 57GE. cf. mithracoides (Frech, 1887) . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . 16, 24, 82, 170, 185; Text-figs 57F, 63E. peracutum (Hall, 1876) . . . . . . . . . . . . 74, 79, 169-170, 171,

185; Text-figs 23, 57A-E; Table 28; Pl. 34, Figs 5, 8E. aff. peracutum (Hall, 1876) . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . 16, 169-171; Text-figs 57B-CE. cf. peracutum (Hall, 1876) . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . 16, 170-171, 185; Text-figs 57D-E, 63equicostatum, Schindewolfoceras? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . 50, 79, 84, 96-97, 98, 209; Text-fig. 23; Pl. 3, Fig. 4equicostatum, Schindewolfoceras? aff. . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . 50, 79, 84, 96-98, 209; Text-fig. 23; Pl. 3, Fig. 1evolutum,Manticoceras cf. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 102Exotornoceras Becker, 1993 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 171expectatum, Clauseniceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 85falcata, Goniatites (Tornoceras) subundulatus var. . . . . . . . . . . . . . . . 173Falcitornoceratini Becker, 1993 . . . . . . . . . . . . . . . . . . . . . . . . . . . . 172FalcitornocerasHouse & Price, 1985 . . . . . . . . . . . . . . . . 73, 77, 171, 173

F. aff. bilobatum (Wedekind, 1908) . . . . . . 172; Text-fig. 58Dfasciculatum, Manticoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 120fasciculatus, Koenenites? . . . . . . . . . . . . . . . . . . . . 1, 27, 29-30, 82, 112-

113, 115, 120-121, 238; Text-figs 33 I-J; Pl. 18, Figs 1-2, 5fasciculatus, Koenenites? cf. . . . . . . . . . . . . . . . . . . . . . . . 31-32, 113, 121FoorditesWedekind, 1918 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 76, 163forcipifer, Acanthoclymenia (G. & F. Sandberger, 1850) . . . . . . . . . . 102frechi, Tornoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 173galeatum, Carinoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . 85, 137-138, 140galeatum, Pharciceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 89, 170Gattendorfia Schindewolf, 1920 . . . . . . . . . . . . . . . . . . . . . . . . . . . 73, 92genundewa, Acanthoclymenia . . . . . . . 1, 27-29, 82, 104-105, 106-107,

155-250; Text-figs 30A-C, K, L; Table 6; Pl. 24, Figs 1-4genundewa, Acanthoclymenia aff. . . . . . . . . . . . . . . . . . . . . . . . . . . . 106genundewa, Acanthoclymenia cf. . . . . . . . . . . . . . . . . . . . 30, 83, 106-107genundewa, Probeloceras cf. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 106?Genus edwinhalli (Clarke, 1898) . . . . . . . . . . . 183, 273; Pl. 35, Fig. 12Gephuroceras Hyatt, 1884 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 123Gephuroceratidae Frech, 1897 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . 72-73, 76-77, 98, 103, 114, 137; Text-figs 20-21Gephuroceratinae Frech, 1879 . . . . . . . . . . . . . . . . . . . . . . . . . . . 122Gephyroceras Carus, 1884 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 123

G. cataphractum Clarke, 1898 . . . . . . . . . . . . . . . . . . . . 141, 153G. holzapfeli Clarke, 1898 . . . . . . . . . . . . . . . . . . . . . 105, 253

gerassimovi, Triainoceras . . . . . . . . . . . . . . . . . . . . . 92, 94; Text-fig. 27DGogoceras Becker et al., 1993 . . . . . . . . . . . . . . . . 73, 77, 98; Text-fig. 23Goniatites acutolateralis G. & F. Sandberger, 1850 . . . . . . . . . . . . . . 182Goniatites affinis Steininger, 1849 . . . . . . . . . . . . . . . . . . . . . . . . . . 142Goniatites astarte Clarke, 1885 . . . . . . . . . . . . . . . . . . . . . . . . . . . . 163Goniatites auris Quenstedt, 1846 . . . . . . . . . . . . . . . . . . . . . . . 175-176Goniatites ausavensis Steiniger, 1853 . . . . . . . . . . . . . . . . . . . . . . . 168Goniatites chemungense Clarke, 1885 . . . . . . . . . . . . . . . . . . . . . . . 93Goniatites cinctumMünster, 1843 . . . . . . . . . . . . . . . . . . . . . . . . . . 176Goniatites complanatus perlatusHall, 1874 . . . . . . . . . . . . . . . . . . 99, 108Goniatites complanatus perlatus Hall, 1876 . . . . . . . . . . . . . . . . . . . . 108Goniatites costatus d’Archiac & de Verneuil, 1842 . . . . . . . . . . . . . . 92Goniatites lamed var. complanatus G. & F. Sandberger, 1850 . . . 132, 136Goniatites lamed var. latidorsalis G. & F. Sandberger, 1850 . . . . . . . 142Goniatites lamed var. tripartitus G. & F. Sandberger, 1850 . . . . . . . . 141

Goniatites lamellosus G. & F. Sandberger, 1850 . . . . . . . . . . . . . . . . 112Goniatites lingua G. & F. Sandberger, 1851 . . . . . . . . . . . . . . . . . . . 116Goniatites lutheri Clarke, 1885 . . . . . . . . . . . . . . . . . . . . . . . . . . . . 156Goniatites mithracoides Frech, 1887 . . . . . . . . . . . . . . . . . . . . . . . . . 169Goniatites nundaia Hall, 1874 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 107Goniatites pattersoni Hall, 1860 . . . . . . . . . . . . . . . . . . . . . . . . . 108Goniatites retrorsus var. lingua G. & F. Sandberger, 1851 . . . . . . . . 166Goniatites retrorsus var. typus G. & F. Sandberger, 1851 . . . . . . . . 163Goniatites simulator Hall, 1874 . . . . . . . . . . . . . . . . . . . . . . . . . . . 122Goniatites sinuosus Hall, 1843 . . . . . . . . . . . . . . . . . . . 107-108, 124Goniatites subpartitusMünster, 1839 . . . . . . . . . . . . . . . . . . . . . . . . 181Goniatites (Tornoceras) subundulatus var. falcata Frech, 1887 . . . . . . 173Goniatites tuberculosocostatus G. & F. Sandberger, 1850 . . . . . . . . . . . 93Goniatites undulatus Brown, 1841 . . . . . . . . . . . . . . . . . . . . . . . . . . 175Goniatites undulatus G. & F. Sandberger, 1851 . . . . . . . . . . . . . . . . 179Goniatites uniangulare Conrad, 1842 . . . . . . . . . . . . . . . . . . 162-163, 164guestphalicum, Tornoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 166Gundolficeras Becker, 1995 . . . . . . . . . . . . . . . . . . . . . . . . . . 73, 162, 173hassoni, Lobotornoceras . . . . . . . . . . . . . . . . . . . 171-172; Text-fig. 58Ehassoni, Lobotornoceras aff. . . . . . . . . . . . . . . . . . . . . . 31-32, 79, 83, 171-

172, 270; Text-figs 23, 58A-B; Table 30; Pl. 34, Figs 1-4haugi, Tornoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 166Hoeninghausia Gürich, 1896 . . . . . . . 73, 83, 112, 120-122; Text-fig. 21

H. archiaci Gürich, 1896 . . . . . . . . . . . . . . . . . . . . . . . . . . 122H. aff. archiaci Gürich, 1896 . . . . . . . . . . . . . . . . . . . . . . . . . 83H. cf. archiaci Gürich, 1896, House et al., 1985 . . . . . . . . 121H. (Koenenites) hoeninghausia von Buch, 1832 . . . . . . . . . . 122

hoeninghausia, Ammonites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 122hoeninghausia, Hoeninghausia (Koenenites) . . . . . . . . . . . . . . . . . . . . 122hoeninghausia, Koenenites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 112-113holzapfeli, Crickites . . . . . . . . . . 77, 86, 149-150, 155; Text-figs 48D-Fholzapfeli, Gephyroceras . . . . . . . . . . . . . . . . . . . . . . . . . . 105, 153, 253housei, Naplesites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 160-161hunanense,Manticoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 142incertum, Trianoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 92inflexum,Maeneceras . . . . . . . . . . . . . . . . . . . . 87, 182-183; Text-fig. 61Fintumescens,Manticoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 129inversum,Manticoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 137isolatum, ?Crassotornoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 168iynx, Naplesites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79, 83, 156, 160-

161, 185-186, 253; Text-figs 23, 52A-E, 63; Pl. 25, Figs 2-5iynx, Naplesites cf. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 160-161kayseri, Ponticeras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 98keyserlingi, Aulatornoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 176keyserlingi, Ponticeras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 98kilfoylei n. ssp., Koenenites styliophilus . . . . . . . . . . . . . . . . . . . . . . . . . . 1,

27-29, 79, 82, 112, 115, 117, 119-120, 185, 218; Text-figs 23,33F, G, N-O, 34G, 63; Table 10; Pl. 8, Figs 5, 7-8, 12-13

kirchgasseri, Koenenites lamellosus . . . . . . . . . . . . . . . . . . . . . 112-113, 117Kirsoceras Bogoslovsky, 1971 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73, 162koeneni, Costamanticoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 137Koenenitidae Becker & House, 1993 . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . 72-73, 77, 98, 112, 114; Text-figs 20-21KoenenitesWedekind, 1913 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . 3, 13, 17-19, 21, 23, 25, 28-30, 32, 65-66, 73, 77, 82,102, 107, 112-113, 115, 117, 120-123, 129, 166; Text-fig. 21K. beckeri n. sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . 1, 31-34, 79, 83, 112-114, 117, 121-122, 185218;Text-figs 23, 33A-E, 34H, K, 63; Pl. 8, Figs 10-11, 14

K. beckeri n. sp., form D . . . . . . . 115, 122; Text-figs 33E, 34KK. cooperiMiller, 1938 . . . . . . . . . . 112-113, 117, 120; Table 9K.? fasciculatus (Clarke, 1898) . . 1, 27, 29, 79, 82, 112-113, 115,

120-121, 238; Text-figs 23, 33 I-J; Pl. 18, Figs 1-2, 5


K.? cf. fasciculatus (Clarke, 1898) . . . . . . . . 31-32, 82, 113, 121K. hoeninghausia (von Buch, 1832) . . . . . . . . . . . . 112-113, 122K. lamellosus (G. & F. Sandberger, 1850) . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . 112-113, 117, 120-121, 122K. lamellosus kirchgasseri House, 1978 . . . . . . . . . 112-113, 117K. aff. lamellosus (G. & F. Sandberger, 1850) sp. D House &

Kirchgaser, 1993 . . . . . . . . . . . . . . . . . . . . . . 83, 121-122K. cf. lamellosus (G. & F. Sandberger, 1850) . . . . 113, 121-122K. styliophilus (Clarke, 1898) . . . . . . . . . . . . . . . . 22, 26, 112K. aff. styliophilus House & Kirchgasser, 1993 . . . . . . . . . . 119K. styliophilus kilfoylei n. ssp. . . . . . . . . . . . . . . . . . 1, 27-29, 79,

82, 112, 115, 117, 119-120, 185, 218; Text-figs 23, 33F,G, N-O, 34G, 63; Table 10; Pl. 8, Figs 5, 7-8, 12-13

K. styliophilus styliophilus (Clarke, 1898) . . . . 1, 17, 19-20, 22-23, 79, 82, 112-113, 115-117, 119-120, 185, 218; Text-figs 23, 34A-F, I- J, L, 35A, 63; Table 8; Pl. 8, Figs 1-9

K. styliophilus styliophilus (Clarke, 1898) morphotype B . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 115, 117, 120

K. sp. House, 1978 . . . . . . . . . . . . . . . . . . . . . . . . . . 113, 117K. sp. C Kirchgasser & House, 1981 . . . . . . . . . . . . . . . . . 121K. sp. D Kirchgasser & House, 1981 . . . . . . . . . . . . . . . . . 122

Komioceras Bogoslovsky, 1958 . . . . . . . . . . . . . . . . 73, 112; Text-fig. 21kwangsiense, Manticoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 142kweipingense,Manticoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 142lamed,Manticoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . 27, 43, 48-49, 54-55, 57, 79, 85, 132, 134-135, 234;Text-figs 23, 41A-C, 42E; Table 14; Pl. 16, Figs 1-2, 9-10

lamed,Manticoceras aff. . . . . . . . . . . . . . . . . . . . . . . . 54, 57, 79, 85, 134-136, 234; Text-figs 23, 41D-E; Table 15; Pl. 16, Figs 3-8

lamed aff. cordatum, Manticoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . 31, 79, 83, 115, 136; Text-figs 23, 33K-L

lamed cf. cordatum, Manticoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 83lamellosus, Koenenites aff. . . . . . . . . . . . . . . . . . . . . . . . . . 83, 121-122lamellosus, Koenenites aff., sp. D . . . . . . . . . . . . . . . . . . . . . . . . . . . . 122lamellosus, Koenenites cf. . . . . . . . . . . . . . . . . . . . . . . . . . . 113, 121-122lamellosus, Goniatites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 112lamellosus, Koenenites . . . . . . . . . . . . . . . . . . 112-113, 117, 120-121, 122latidorsalis, Goniatites lamed var. . . . . . . . . . . . . . . . . . . . . . . . . . . . 142lepiferum, Aulatornoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 175lindneri, Crickites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . 58, 79, 85, 149-150, 151, 185; Text-figs 23, 49; Table 21lindneri,Manticoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 150lingua, Goniatites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 166lingua, Goniatites retrorsus var. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 166LinguatornocerasHouse, 1965 . . . . . . . . . 73, 162, 166, 168; Text-fig. 21

L. clausum (Glenister, 1958) . . . . . . . . . . . . . . . . . . . . . . . 167L. linguum (G. & F. Sandberger, 1851) . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . 167, 257; Pl. 27, Figs 13-14L. aff. linguum (G. & F. Sandberger, 1851) . . . . . . . . . . . . . . . .

. . . 43, 48, 54-55, 79, 84-85, 167, 257, 269; Text-figs 23,56A-F; Table 29; Pl. 27, Fig. 12, Pl. 33, Figs 1-2, 5-8

L. sp. . . . . . . . . . . . . . . . . . . . . . . . 167, 269; Pl. 33, Figs 3-4linguum, Linguatornoceras . . . . . . . . . . . . . 167, 257; Pl. 27, Figs 13-14linguum, Linguatornoceras aff. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . 43, 48, 54-55, 79, 84-85, 167, 257, 269; Text-figs23, 56A-F; Table 29; Pl. 27, Fig. 12, Pl. 33, Figs 1-2, 5-8

linguum, Tornoceras (Linguatornoceras) aff. . . . . . . . . . . . . . . . . . . . 166Lobotornoceras Schindewolf, 1936 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . 65, 73, 162, 168, 171-172; Text-fig. 21L. ausavense (Steininger, 1853) . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . 168, 171-172; Text-figs 58C, FL. bicaniculatum Petter, 1959 . . . . . . . . . . . . . . . . . . . . . . . . 173L. hassoni House, 1978 . . . . . . . . . . . . . . . 172; Text-fig. 58E

L. aff. hassoni . . . . . . . . . . . . . . . . . . . . . . . . 31-32, 79, 83, 172,270; Text-figs 23, 58A-B; Table 30; Pl. 34, Figs 1-4

loeschmanni, Aulatornoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . 169, 176lutheri, Probeloceras aff. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 157lutheri, Probelcoeras cf. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 157, 160lutheri, Goniatites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 156lutheri, Probeloceras . . . . . . . . . . . . . . . . . . . . . . . . 35, 38-43, 79, 95, 105,

125, 152, 155-156, 157, 159-161, 185, 250, 253; Text-figs 23,35C, 50-51, 63; Table 22; Pl. 24, Figs 5-9, Pl. 25, Fig. 1

Maeneceras Hyatt, 1844 . . . . . . . . 62, 73, 87, 182-183; Text-figs 21, 61M. acutolaterale (G. & F. Sandberger, 1850) . . . . . . 86, 183, 185M. aff. acutolaterale (G. & F. Sandberger, 1850) . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . 62, 79, 86, 182-183, 184-185,273; Text-figs 23, 61C-D, 62-63; Pl. 35, Figs 11, 13

M. inflexum (Wedekind, 1908) . . . . 87, 182-183; Text-fig. 61FM. milleri (Flower & Caster, 1935) . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . 3, 62, 185; Text-figs 61E, 63M. pompeckji (Wedekind, 1918) . . . . . . . . . . . . . . . . . . . . . 86M. cf. pompeckji (Wedekind, 1918) . . . . . . . . . . . . . . . 62, 183

Maenioceras Schindewolf, 1933 . . . . . . . . . . . . . . . . . . . . . . . . 76, 81, 90Maenioceratidae Bogoslovsky, 1958 . . . . 71-72, 73, 76; Text-figs 20-21Manticoceratidae Wedekind, 1913 . . . . . . . . . . . . . . . . . . . . . . . . . 122Manticoceras Hyatt, 1884 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . 2, 43, 51, 73, 76-77, 83, 102-103, 105, 107, 112-113,115, 120, 122-124, 129, 135, 137, 142, 150, 157; Text-fig. 21M. accelerans Clarke, 1887. . . . . . . . . . . . . . . . . . . . . 1, 137, 237M. altaicum Bogoslovsky, 1958 . . . . . . . . . . . . . . . . . . . . . 150M. apprimatum Clarke, 1898 . . . . . . . . . . . . . . . . . . . 123, 132M. buchii (d’Archiac & de Verneuil, 1842) . . . . . . . . . . . . 129M. contractum Clarke, 1898 . . 1, 27, 29, 79, 83, 118, 123, 130,

132, 136-137, 241; Text-figs 23, 35B; Pl. 19, Figs 10-16M. cordatum (G. & F. Sandberger, 1850) . . 129, 134, 136M. cordiformeMiller, 1938 . . . . . . . . . . . . . . . . . . . . 129, 150M. evolutum Petter, 1959 . . . . . . . . . . . . . . . . . . . . . . . . . . 102M. cf. evolutum House & Kirchgasser, 1993 . . . . . . . . . . . 102M. fasiculatum Clarke, 1898 . . . . . . . . . . . . . . . . . . . . . . . 120M. hunanense Xu, 1977 . . . . . . . . . . . . . . . . . . . . . . . . . . . 142M. intumescens (Beyrich, 1837) . . . . . . . . . . . . . . . . . . . . . 129M. inversumWedekind, 1913 . . . . . . . . . . . . . . . . . . . . . . 137M. kwangsiense Chao, 1956 . . . . . . . . . . . . . . . . . . . . . . . . 142M. kweipingense Chao, 1956 . . . . . . . . . . . . . . . . . . . . . . . 142M. lamed (G. & F. Sandberger, 1850) . . . . . . . . . . . . . . . . . . . .

27, 43, 48-49, 54-55, 57, 79, 85, 132, 134-135, 234; Text-figs 23, 41A-C, 42E; Table 14; Pl. 16, Figs 1-2, 9-10

M. lamed aff. cordatum (G. & F. Sandberger, 1850) . . . . . . . . . .. . . . . . . . . . . . 31, 79, 83, 115, 136; Text-figs 23, 33K-L

M. aff. lamed (G. & F. Sandberger, 1850) . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . 54, 57, 79, 85, 134-136,234; Text-figs 23, 41D-E; Table 15; Pl. 16, Figs 3-8

M. lindneri Glenister, 1958 . . . . . . . . . . . . . . . . . . . . . . . . 150M. neverovi Bogoslovsky, 1958 . . . . . . . . . . . . . . . . . . . . . . 150M. nodifer (Clarke, 1885) . . . . . . . . . . . . . . . . . . . . . . . . . . 27,

79, 83, 123, 238; Text-figs 23, 35B; Pl. 18, Figs 11-12M. oxy Clarke, 1897 . . . . . . . . . . . . . . . . . . . . . . . . 55, 137, 142M. Pattersoni Clarke, 1898 . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . 113, 115, 117, 119, 124-125, 127, 129-132, 221, 230M. pattersoni var. styliophilum Clarke, 1898 . . . . . 113, 115, 119M. regulare Fenton & Fenton, 1924 . . . . . . . . . . . . . . . . . . 129M. rhynchostomum Clarke, 1898 . . . . . . . . . . . . . . . . . . . . 142M. schellwieniWedekind, 1913 . . . . . . . . . . . . . . . . . . . . . 137M. septentrionaleMiller, 1938 . . . . . . . . . . . . . . . . . . 129, 150M. simulator (Hall, 1874) . . 83, 124, 130, 221; Pl. 9, Figs 4-5M. sinuosum (Hall, 1843) . . . . . . . . . . . . . . . . . . . . . . . . . . . . .


. . . . 1, 34, 38-42, 108, 115, 123, 129, 131-132, 135, 144M. cf. sinuosum House & Pedder, 1963 . . . . . . . . . . . . . . . 129M. sinuosum apprimatum Clarke, 1898 . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . 1, 27, 79, 83, 123, 132, 135,185, 237; Text-figs 23, 35B, 42F, 63; Pl. 17, Figs 5-10

M. sinuosum clausium n. ssp. . . . . . . . . . . . . . . . . . . . . . . . . . 1,35, 40-41, 79, 129-131, 226, 229; Text-figs 23, 39A-I,40A-B; Table 12; Pl. 12, Figs 1-7, 11, Pl. 13, Figs 1-15

M. sinuosum sinuosum (Hall, 1843) . . . . . . . . . . . . . . . . . 35, 38-42, 79, 83, 124-127, 129-130, 132, 135, 221-222, 225-226, 233-234; Text-figs 23, 36A-G, 37A-H, 38; Table 11;Pl. 9, Figs 6-8, Pl. 10, Figs 1-12, Pl. 11, Figs 1-13, Pl. 12,Figs 8-10; Pl. 15, Figs 1-2, Pl. 16, Figs 11-12

M. sinuosum tardum Clarke, 1898 . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . 1, 40, 79, 130, 132, 221, 230; Text-figs 23,39J, 40C; Table 13; Pl. 9, Figs 2-3, Pl. 14, Figs 1-10

M. tardum Clarke, 1898 . . . . . . . . . . . . . . . . . . . . . . . 130-132M. unduloconstrictumMiller, 1938 . . . . . . . . . . . . . . . . . . . 141M. sp. . . . . . . . . . . . . . . . . . . . . . . . . 137, 237; Pl. 17, Fig. 4

manticoides, Chutoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 107-109Maternoceras Clausen, 1971 . . . . . . . . . . . . . 73, 77, 122; Text-fig. 21

M. sandbergeri (Wedekind, 1910) . . . . . . . . . . . . . . . . . . . . . 85menneri, Carinoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 137-138Meropharciceras Becker & House, 1993 . . . . . . . . . . 73, 76; Text-fig. 21Mesobeloceras Glenister, 1958 . . . 73, 77, 83, 155, 160-161; Text-fig. 21milleri,Maeneceras . . . . . . . . . . . . . . . . . . . 3, 62, 185; Text-figs 61E, 63milleri, Sporadoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 62mithracoides, Epitornoceras . . . . . . . . . . . . . . . . 82, 169-170; Text-fig. 57Gmithracoides, Epitornoceras cf. . . . 16, 24, 82, 170, 185; Text-figs 57F, 63mithracoides, Goniatites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 169Mixomanticoceras Becker et al., 1993 . . . . . . . . . . . . . . 122; Text-fig. 21Mzerrebites Becker & House, 1993 . . . . . . . . . . . . . . 73, 76; Text-fig. 21naplesense, Naplesites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . 79, 160-161, 186, 253; Text-figs 23, 52F-H; Pl. 25, Fig. 6naplesense, Neomanticoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 160naplesense, Probeloceras? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 160, 253Naplesites Yatskov, 1991 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . 4, 73, 77, 83, 155-157, 160-161, 186, 253; Text-figs 21, 52N. anguisellatum (Chao, 1956) . . . . . . . . . . . . . . . . . 160-161N. housei (Montesinos & Henn, 1986) . . . . . . . . . . . . . . . 161N. inyx (Clarke, 1898) . . . . . . . . . . . . . . . . . . . . . . 79, 160-161,

185-186, 253; Text-figs 23, 52A-E, 63; Pl. 25, Figs 2-5N. cf. inyx Becker, 1993 . . . . . . . . . . . . . . . . . . . . . . . . . . 161N. naplesense (Clarke, 1898) . . . . . . . . . . . . . . . . . . . . . . . . . 79,

160-1671, 186, 253; Text-figs 23, 52F-H; Pl. 25, Fig. 6N. sp. (Bogoslovsky, et al., 1982) . . . . . . . . . . . . . . . . . . . . 160

neapolitana, Acanthoclymenia . . . . . . . . . . . . . . . 35, 39-40, 79, 83, 103-106, 214; Text-figs 23, 30E-H; Table 5; Pl. 6, Figs 1-15

neapolitana, Acanthoclymenia aff. . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . 31, 33, 79, 83, 102-105; Text-figs 23, 30D

neapolitana, Acanthoclymenia cf. . . . . . . . . . . . . . . . . . . . . . . . . . . . 106Neomanticoceras Schindewolf, 1936 . . 73, 84-85, 122, 160; Text-fig. 21

N. naplesense (Clarke, 1898) . . . . . . . . . . . . . . . . . . . . . . . 160N. paradoxum (Matern, 1931) . . . . . . . . . 217; Pl. 7, Figs 2-3

Neopharciceras Bogoslovsky 1955 . . . . . . . 73, 76-77, 82, 89; Text-fig. 21neverovi,Manticoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 150nitidum, Crassotornoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 168nodifer, Manticoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

27, 79, 83, 123, 238; Text-figs 23, 35B; Pl. 18, Figs 11-12nundaia, Goniatites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 107nundaium, Chutoceras . . . . . . . . . . . . . . 1, 17, 23, 25-26, 77, 79, 82, 100,

107-109, 185, 221; Text-figs 23, 28H-K, 63; Pl. 9, Fig. 1obseum, Tornoceras uniangulare . . . . . . . . . . . . . . . . 35, 39-41, 79, 83, 158,

165, 258; Text-figs 23, 54G, 55C; Table 25; Pl. 28, Figs 6-10orbiculus, Ammonites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 142orientale, Acanthoclymenia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 102ovata, Tornoceras simplex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 167oxy, Carinoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 144oxy, Manticoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55, 137, 142oxy, Sphaeromanticoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . 1, 43, 45, 48, 54-55, 57-58, 79, 85, 135, 140, 143-144, 145,148; Text-figs 23, 42D, 44A, C, E-F, 45A-D, 46B; Table 19

oxy, Sphaeromanticoceras aff. . . . . . . . . 43, 48, 85, 140, 148; Text-fig. 44FOxytornoceras Becker, 1993 . . . . . . . 73, 77, 86, 162, 169; Text-fig. 21

O. acutum (Frech, 1902) . . . . . . . . . . . . . . . . . . . . . . . . . . 169paradoxum, Neomanticoceras . . . . . . . . . . . . . . . . . . 217; Pl. 7, Figs 2-3Parodiceras Hyatt, 1884 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 162Parodiceratinae Petter, 1959 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 162pattersoni, Goniatites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 108Pattersoni,Manticoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . 113, 115, 117, 119, 124-125, 127, 129-132, 221, 230paucistriatum, Aulatornoceras . . . . . . . . . . 54-55, 79, 85, 175-176, 178-

180, 266; Text-figs 23, 60D, F; Table 33; Pl. 32, Figs 2-13peracutum, Epitornoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24, 79, 169-

170, 171, 185; Text-figs 23, 57A-E; Table 28; Pl. 34, Fig. 8peracutum, Epitornoceras aff. . . . . . . . . . 16, 169-171; Text-figs 57B-Cperacutum, Epitornoceras cf. . . . . 16, 170-171, 185; Text-figs 57D-E, 63perlatum, Ponticeras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16-17, 20,

24-26, 79, 82, 98-99, 100-102, 108, 185, 210, 213; Text-figs28A-G, 29A-E, 63; Table 4; Pl. 4, Figs 5-9, Pl. 5, Figs 1-9

perlatum, Ponticeras cf. . . . . . . . . . . . 16, 23, 101-102; Text-figs 29D-EPetteroceras Bogoslovsky, 1967 . . . . . . . . . . . . . . 73, 76, 82; Text-fig. 21Petteroceratidae Becker & House, 1993 . . . 72-73, 89; Text-figs 20-21Pharciceras Hyatt, 1884 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . 3, 10-11, 66, 73, 76, 89-92, 186, 205; Text-fig. 21P. amplexum (Hall, 1886) . . 10-11, 79, 82, 89-90, 91, 185-186,

205; Text-figs 23, 26D-H, 63; Table 2; Pl. 1, Figs 1-8P. barnettiWork, Mason, & Klapper, 2007. . . . . . . . . . . . . . . 186P. bidentatum Petter, 1959 . . . . . . . . . . . . . . . . . . . . . . . . . . . 89P. galeatumWedekind, 1918 . . . . . . . . . . . . . . . . . . . . 89, 170P. tridens (G. & F. Sandberger, 1850) . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . 1, 89-91, 205; Text-figs 26 A-C; Pl. 1, Fig. 10P.? sp. . . 13, 27, 79, 92, 265; Text-fig. 23; Pl. 31, Figs 14-15

Pharciceratidae Hyatt, 1900 . . . . . . . . . . . . . 72-73, 89; Text-figs 20-21Phoenixites Becker, 1993 . . . . . . . . . . . . . . 60, 73, 77, 173; Text-fig. 21

P. concentricus (House, 1965) . . . 59-60, 79, 86, 158, 173, 261;Text-figs 23, 54M-O, 56A; Table 26; Pl. 29, Figs 1-15

P. sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 62planorbe, Acanthoclymenia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 102Playfordites Becker et al., 1993 . . . 73, 84-85, 122, 135, 141; Text-fig. 21

P. tripartitus (G. & F. Sandberger, 1850) . . . . . . . . . . . . . . 141P. cf. tripartitus (G. & F. Sandberger, 1850) . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . 43, 48, 54, 57, 79, 84-85, 135, 142, 185; Text-figs 23, 42A, 63; Table 17

pompeckji, Maeneceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86pompeckji,Maeneceras cf. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 62, 86pompeckji, Sporadoceras cf. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 183pompeckji, Tornoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 166PonticerasMatern, 1929 . . . . . . . . . . . . . . . . . . . . . . . . . . . 4, 13, 16, 19-

21, 23, 27, 66, 73, 77, 82, 98, 100-103, 107-109; Text-fig. 21P. aequabile (Beyrich, 1837) . . . . . . . . . . . . . . . . . . . . . . . . 98P. auritum (Holzapfel, 1899) . . . . . . . . . . . . . . . . . . . . . . . 98P. barroisi (Wedekind, 1917,1918) . . . . . . . . . . . . . . . . . . . 98P. bisulcatum (Keyserling, 1844) . . . . . . . . . . . . . . . . . . . . 98P. discoidale Glenister, 1958 . . . . . . . . . . . . . . . . . . . . . . . . 98P. domanicense (Holzapfel, 1899) . . . . . . . . . . . . . . . . . . . . 98


P. kayseri (Wedekind, 1917, 1918) . . . . . . . . . . . . . . . . . . . 98P. keyserlingi (Holzapfel, 1899) . . . . . . . . . . . . . . . . . . . . . . 98P. perlatum (Hall, 1874, 1876) . . . . . 16-17, 20, 24-26, 79, 82,

98-99, 100-102, 108, 185, 210, 213; Text-figs 23, 28A-G,29A-E, 63; Table 4; Pl. 4, Figs 5-9, Pl. 5, Figs 1-9

P. cf. perlatum (Hall, 1874) . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . 16, 23, 101-102; Text-figs 29D-E

P. cf. regale (Holzapfel, 1899) . . . . . . . . . . . . . . . . 82, 107-108P.? sp. . . . . . . . . . . . . . . 31, 79, 83, 102, 115; Text-figs 23, 33M

posterior, Aulatornoceras eifliense . . . . . . . . . . . . . . . . . . . . . . . . . . . . 175Posttornoceratidae Bogoslovsky, 1962 . . . . . . . 72, 161-162; Text-fig. 20primordalis, Ammonites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 142Probeloceras Clarke, 1898 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4, 35,

73, 82, 102-103, 105, 109, 155-156, 157, 160; Text-fig. 21P. alveolatum Glenister, 1958 . . . . . . . . . . . . . . . . . . . . . . . 109P. cf. genundewa Clarke, 1898; House, 1978 . . . . . . . . . . . 106P. lutheri (Clarke, 1855) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . 35, 38-43, 79, 95, 105, 125, 152, 155-156, 157, 159-161, 185, 250, 253; Text-figs 23, 35C, 50-51, 63; Table 22; Pl. 24, Figs 5-9, Pl. 25, Fig. 1

P. lutheri n. ssp. Becker et al., 1993 . . . . . . . . . . . . . . . . . . 157P. aff. lutheri Becker et al., 1993 . . . . . . . . . . . . . . . . . . . . . 157P. cf. lutheri Becker et al., 1993 . . . . . . . . . . . . . . . . . . . 157, 160P. strix Kirchgasser, 1975 . . . . . . . . . . . . . . . . . . . . . . . . 83, 109P. aff. strix Kirchgasser, 1975 . . . . . . . . . . . . . . . . . . . . . . . 110P.? naplesense Clarke, 1898 . . . . . . . . . . . . . . . . . . . . . . . 160, 253

Prochorites Clausen, 1969 . . . . 73, 98, 102, 105, 109, 157; Text-fig. 21P. alveolatus (Glenister, 1958) . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . 35, 38-41, 79, 83, 109-110, 156-157, 185,217; Text-figs 23, 31, 32C, 63; Table 7; Pl. 7, Figs 4-6

P. aff. alveolatus (Glenister, 1958) . . . . 49, 79, 110, Text-fig. 23Prolecanites Majsisovics, 1882 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 97Pseudarietites Frech, 1902 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 92Pseudoclymeniidae sensu Becker, 1995 . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . 72-73, 78, 161-162; Text-figs 20-21Pseudofordites Bogoslovsky, 1959 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 76Pseudoprobeloceras Bensaïd, 1974 . . . . . 73, 76, 98, 120, 122; Text-fig. 21Puncticeras Becker, 1993 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 181Raymondiceras Schindewolf 1934 . . . . . . . . . . . . . . . . . . . . . . . . . . 181regale, Ponticeras cf. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 82, 107-108regulare, Manticoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 129rhynchostomum, Manticoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 142rhynchostomum, Sphaeromanticoceras . . . . . . . . . . . . . . . . . . 1, 43, 48-49,

51, 54-57, 79, 84-85, 134-135, 139, 142-144, 148, 185, 242,245-246; Text-figs 23, 41F, 42B, H-I, 43F-G, 47, 63; Table 18;Pl. 20, Figs 1-12, Pl. 21, Figs 1-12, Pl. 22, Figs 1-9

rhysum, Aulatornoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 58-59, 79,85, 175-176, 178-179, 266; Text-figs 23, 60E; Pl. 32, Fig. 1

rickardi, Sphaeromanticoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 58-59, 79, 148-149, 155, 185; Text-figs 23, 48A-C, 63; Table 20

rickardi, Sphaeromanticoceras aff. . . . . . 58, 79, 85, 149-150; Text-fig. 23sagittarium, Beloceras . . . . . . . . . . . . . . . . . . . . . . . . . . 94; Text-fig. 27Csandbergeri, Aulatornoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 175sandbergeri, Maternoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 85sandbergeri, Tornoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 179sandbergeri, Truyolsoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 179Sandbergeroceras Hyatt, 1884 . . . 49-50, 73, 83, 92-93, 95; Text-fig. 21

S. Chemungense Clarke 1898 . . . . . . . . . . . . . . . . . . . . . . . 97S.? enfieldense n. sp. . . . . . . . . . . . . . . . . . . 1, 49-51, 79, 95,

206, 209; Text-fig. 23; Pl. 2, Figs 1-2, Pl. 3, Figs 2-3S. sandbergerorum Miller, 1938 . . . . . . . . 92-95; Text-fig. 27FS. syngonum (Clarke, 1897) . . . . . . . . . . . . . . . . . . . . . . . . . . . .

1, 34, 42, 49-50, 79, 93-95, 185, 206, 209; Text-figs 23,

27G-H, 63; Table 3; Pl. 2, Figs 4-7, Pl. 3, Figs. 5-6S.? syngonum Miller, 1938 . . . . . . . . . . . . . . . . . . . . . . . . 95

Sandbergeroceratidae Miller, 1938 . . . . . . . . . . . . . . . . . . . . . . . . . . 89sandbergerorum, Sandbergeroceras . . . . . . . . . . . . . 92-95; Text-fig. 27Fschellwieni, Manticoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 137SchindewolfocerasMiller, 1938 . . . . . 49, 73, 77, 84, 92, 96; Text-fig. 21

S. chemungense (Vanuxem, 1842) . . . . . . . . . . . . . . . . 50, 79, 84,94, 96-97, 98, 206; Text-figs 23, 27I, 63; Pl. 2, Fig. 8

S. aff. chemungense (Vanuxem, 1842) . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . 94, 97, 210; Text-fig. 27J; Pl. 4, Figs 1-4

S.? equicostatum (Hall, 1874) . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . 50, 79, 84, 96-97, 98, 209; Text-fg. 23; Pl. 3, Fig. 4

S.? aff. equicostatum (Hall, 1874) . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . 50, 79, 84, 96-98, 209; Text-fig. 23; Pl. 3, Fig. 1

?Schindewolfoceras sp. . . . . . . . . . . . . . . . . . . . 51, 97, 206; Pl. 2, Fig. 3Semicheiloceras Becker, 1993 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 162septentrionale,Manticoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . 129, 150Serramanticoceras Becker, House, & Kirchgasser, 1992 . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73, 77; Text-fig. 21serriense, Aulatornoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 176simplex, Ammonites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 163simplex, Tornoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 163simulator, Goniatites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 122-124simulator, Manticoceras . . . . . . . . . . . 83, 124, 130, 221; Pl. 9, Figs 4-5sinuosum,Manticoceras cf. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 129sinuosum, Manticoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . 1, 34, 38-42, 108, 115, 123, 129, 131-132, 135, 144sinuosum sinuosum, Manticoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . 35, 38-42, 79, 83, 124-127, 129-130, 132,135, 221, 225-226, 233-234; Text-figs 23, 36A-G, 37A-H, 38;Table 11; Pl. 9, Figs 6-8, Pl. 10, Figs. 1-12, Pl. 11, Figs 1-13,Pl. 12, Figs 8-10, Pl. 15, Figs 1-2, Pl. 16, Figs 11-12

sinuosus, Goniatites . . . . . . . . . . . . . . . . . . . . . . . . . . . . 107-108, 124sororium, Carinoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . 79, 85, 137, 238; Text-figs 23, 42G; Pl. 18, Figs 3-4, 6-9Sphaeromanticoceras Clausen, 1971 . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

57, 73, 77, 122, 129, 134-135, 137, 140, 142, 149; Text-fig. 21S. affine (Steininger, 1849) . . . . . . . . . . . . . . . . . . . . . . 142, 157S. oxy (Clarke, 1897) . . . . . . . . . . . . . . . . . . . . . . . . 1, 43, 45,

48, 54-55, 57-58, 79, 85, 135, 140, 143-144, 145, 148;Text-figs 23, 42D, 44A, C, E, 45A-D, 46B; Table 19

S. aff. oxy (Clarke, 1897) . . . 43, 48, 85, 140, 148; Text-fig. 44FS. rickardiHouse & Kirchgasser, 1993 . . . . . . . . . . . . 58-59, 79,

148-149, 155, 185; Text-figs 23, 48A-C, 63; Table 20S. aff. rickardiHouse & Kirchgasser, 1993 . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . 58, 79, 85, 149-150; Text-fig. 23S. rhynchostomum (Clarke, 1898) . . 1, 43, 48-49, 51, 54-57, 79,

84-85, 134-135, 139, 142-144, 148, 185, 242, 245-246;Text-figs 23, 41F, 42B, H-I, 43F-G, 47, 63; Table 18; Pl.20, Figs 1-12, Pl. 21, Figs 1-12, Pl. 22, Figs 1-9

Sphaeropharciceras Bogoslovsky, 1955 . . . . . . . . . . . . . . . . . . . . . . . 89Sporadoceras Hyatt, 1884 . . . . . . . . . . . 3, 73, 87, 182-183; Text-fig. 21

S. milleri (Flower & Caster, 1935) . . . . . . . . . . . . . . . . . . . 62S. cf. pompeckjiWedekind, 1918 . . . . . . . . . . . . . . . . . . . . 183

Sporadoceratidae Miller & Furnish, 1957 . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . 72-73, 182-183; Text-figs 20-21

StaffitesWedekind, 1918 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77, 181StenopharcicerasMontesinos & Henn, 1986 . . . 73, 76, 89; Text-fig. 21strix, Eidoprobeloceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 109strix, Probeloceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 83, 109strix, Probeloceras aff. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 110styliophilum,Manticoceras pattersoni var. . . . . . . . . . . . . . . . 113, 115, 119styliophilus, Koenenites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22, 26, 112


styliophilus, Koenenites aff. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 119styliophilus styliophilus, Koenenites . . . . . . . . . . . . . . . . . . . . . . . . . . 1, 17,

19-20, 22-23, 79, 82, 112-113, 115-117, 119-120, 185, 218;Text-figs 23, 34A-F, I-J, L, 35A, 63; Table 8; Pl. 8, Figs 1-9

styliophilus styliophilusmorphotype B, Koenenites . . . . . . . . . 115, 117, 120subpartitus, Goniatites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 181syngonum, Sandberberoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . 1, 34, 42, 49-50, 79, 93-95, 185, 206, 209;Text-figs 23, 27G-H, 63; Table 3; Pl. 2, Figs 4-7, Pl. 3, Figs 5-6

syngonum, Sandbergeroceras? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 95Synpharciceras Schindewolf, 1940 . . . . . . . . . . . . 73, 76, 89; Text-fig. 21Synpharciceratidae Schindewolf, 1940 . . . . . . . . . . . . . . . . . . . . . . . . 89Tamarites Bogoslovsky, 1965 . . . . . . . . . . . . . . . . 73, 91-92; Text-fig. 21tardum, Manticoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 130-132tardum,Manticoceras sinuosum . . . . 1, 40, 79, 130, 132, 221, 230; Text-

figs 23, 39J, 40C; Table 13; Pl. 9, Figs 2-3, Pl. 14, Figs 1-10TimanitesMojsisovics, 1882 . . . . . . . . . . . . . . . 73, 83, 112; Text-fig. 21Timanoceras Bogoslovsky, 1957 . . . . . . . . . . . . 73, 77, 122; Text-fig. 21TorniaHouse, 1970 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73, 78, 162TornocerasHyatt, 1884 . . . . . . . . . . 11, 16, 19-21, 27-34, 41-42, 60, 65,

73, 76, 157-158, 162-167, 169, 171, 173; Text-figs 21, 53-54T. acutum Frech, 1902 . . . . . . . . . . . . . . . . . . . . . . . . . . . . 169T. arcuatumHouse, 1965 . . 22, 79, 82, 158, 165-166, 170;Text-

figs 23, 54H-J, L; Pl. 26, Figs 7-10, Pl. 27, Fig. 16T. cf. arcuatum House, 1965 . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . 10, 158, 166, 170; Text-fig. 54KT. ausavense crassumMatern 1931 . . . . . . . . . . . . . . . . . . . 168T. belgicum (Matern, 1931) . . . . . . . . . . . . . . . . . . . . . . . . . 168T. bicostatum (Hall, 1843) . . . . . . . . . . . . . . . . . . . . . . . 178T. (Aulatornoceras) bicostatum (Hall, 1843) . . . . . . . . . . 178-179T. (Aulatornoceras) clarkeiMiller, 1938 . . . . . . . . . . . . . . . . 61T. clausum Glenister, 1958 . . . . . . . . . . . . . . . . . . . . . . . . . 166T. (Tornoceras) concentricum House, 1965 . . . . . . . . . . . . . 173T. Edwin-halli Clarke, 1898 . . . . . . . . . . . . . . . . . . . 62, 87, 183T. eifliense (Steininger, 1849) . . . . . . . . . . . . . . . . . . . . . . . 177T. frechiWedekind, 1918 . . . . . . . . . . . . . . . . . . . . . . . . . . 173T. guestphalicum Frech, 1897 . . . . . . . . . . . . . . . . . . . . . . . 166T. haugi Frech, 1902 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 166T. (Linguatornoceras) aff. linguum House, 1965 . . . . . . . . . 166T. pompeckjiWedekind 1918 . . . . . . . . . . . . . . . . . . . . . . . 166T. sandbergeri (Ford & Crick, 1897) . . . . . . . . . . . . . . . . . 179T. simplex (von Buch, 1832) . . . . . . . . . . . . . . . . . . . . . . 163T. simplex mut. ovata (Münster, 1832) Frech, 1902 . . . . . . 167T. typum (G. & F. Sandberger, 1851) . . . . . . . . . . . . . . . . 16, 163T. cf. typum (G. and F. Sandberger, 1851) . . . . . . . . . 43, 48-49,

55, 85, 157, 166, 257; Text-figs 53C-D; Pl. 27, Fig. 15T. uniangulare (Conrad, 1842) . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . 39, 163, 165-166, 169; Text-fig. 23T. aff. uniangulare (Conrad, 1842) . . . . . . . . . . . . . . . . . . . . . . .

. . . 11, 79, 158, 164, 257; Text-fig. 23; Pl. 27, Figs 1-6T. cf. uniangulare (Conrad, 1842) . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . 79, 158, 164, 185; Text-figs 23, 54D-E, 63T. uniangulare arkonensis (House, 1965) . . . . . . . . . . . . . . . . . 81T. uniangulare compressum Clarke, 1897 . . . . . . . . . . . . . . . . . .

. . . . . . . 27, 29, 79, 82, 157-158, 162, 164-165, 158; Text-figs 23, 53A-B, 54F, 55B; Table 24; Pl. 28, Figs 1-5

T. uniangulare obesum Clarke, 1897 . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . 35, 39-41, 79, 83, 158, 165, 258;Text-figs 23, 54G, 55C; Table 25; Pl. 28, Figs 6-10

T. uniangulare uniangulare (Conrad, 1842) . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . 1, 10-12, 79, 158, 160, 163, 165, 254;Text-figs 23, 54A-C, 55A; Table 23; Pl. 26, Figs 1-4, 6

T. uniangulare widderi House, 1965 . . . 81, 254; Pl. 26, Fig. 5

T. sp. House, 1965 . . . . . . . . . . . . . . . . . . . . . . . Text-fig. 54PT.? n. sp. House, 1965 . . . . . . . . . . . . . . . . 257; Pl. 27, Fig. 11

Tornoceratidae Arthaber, 1911 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . 71-72, 73, 161-162, 172-173, 175; Text-figs 20-21

Tornoceratinae Arthaber, 1911 . . . . . . . . . . . . . . . . . . . . . . . . . . . . 162Tornoceratini Arthaber, 1911 . . . . . . . . . . . . . . . . . . . . . 162, 172-173Trevonites Becker & House, 1994 . . . . . . . . . . . . . . 73, 162, Text-fig. 21Triainocerae Hyatt, 1884 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 89, 92Triainoceras Hyatt, 1884 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 91-93

T. costatum (d’Archiac & de Verneuil, 1842) . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 94; Text-fig. 27E

T. gerassimovi Bogoslovsky, 1958 . . . . . . . . 92, 94; Text-fig. 27DT. incertum (d’Archiac & de Verneuil, 1842) . . . . . . . . . . . 92T.? tuberculosum (d’Archiac & de Verneuil, 1842) . . . . . . . 92

Triainoceratidae Hyatt, 1884 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .1, 50, 63, 71-72, 73, 83-84, 88-89, 91-92, 94; Text-figs 20-21

tridens, Pharciceras . . . . 1, 89-91, 205; Text-figs 26 A-C; Pl. 1, Fig. 10Trimanticoceras House in House & Ziegler, 1977 . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73, 77, 122; Text-fig. 21T. cinctum (Glenister, 1958) . . . . . . . . . . . . . . . . . . . . . . . . 85

tripartitus, Playfordites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 141tripartitus, Playfordites cf. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43, 48, 54,

57, 79, 84-85, 135, 142, 185; Text-figs 23, 42A, 63; Table 17tripartitus, Goniatites lamed var. . . . . . . . . . . . . . . . . . . . . . . . . . . . 141TruyolsocerasMontesinos, 1987 . . . . . . . . . . . 73, 175, 179; Text-fig. 21

T. bicostatum (Hall, 1843) . . . . . . . . . . . . . . . . . . . . . . . . . 1, 59-61, 79, 86, 175-176, 178-179, 180, 257, 262; Text-figs 23,59B, 60A-C, H; Table 34; Pl. 30, Figs 2, 4-13, 15

T. cf. bicostatum (Hall, 1843) . . . . . . . . . . . 59-61, 79, 86, 180,262, 265; Text-fig. 23; Pl. 30, Fig. 15, Pl. 31, Fig. 1

T. clarkei (Miller, 1938) . . . . . . . . . . . . . . . . . . . . . . . . . . 59, 61,79, 86, 176, 179-180, 262; Text-fig. 23; Pl. 30, Figs 1, 3

T. sandbergeri (Ford & Crick, 1897) . . . . . . . . . . . . . . . . . 179tuberculoso-costatus (pro, pars) G. & F. Sandberger, 1850, Goniatites . . . 93tuberculosum, Trianoceras? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 92tynani, Wellsites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50-51, 79,

83-84, 94, 96, 185, 205; Text-figs 23, 27A, 63; Pl. 1, Fig. 9typum, Tornoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16, 163typum, Tornoceras cf. . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . 43, 48-

49, 55, 85, 157, 166, 257; Text-figs 53C-D; Pl. 27, Fig. 15typus, Goniatites retrorsus var. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 163Uchtites Bogoslovsky, 1969 . . . . . . . . . . . . . . . . . . . 73, 98; Text-fig. 21undulatus, Goniatites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 175unduloconstrictum, Delphiceras . . . . . . . . . . . . . . . . . 238; Pl. 18, Fig. 10unduloconstrictum,Manticoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . 141uniangulare, Tornoceras . . . . . . . . . . . . . . . . . . . . . . . . . . Text-fig. 23uniangulare, Tornoceras aff. . . . . . . . . . . . Text-fig. 23; Pl. 27, Figs 1-6uniangulare, Tornoceras cf. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . 79, 158, 164, 185; Text-figs 23, 54D-E, 63uniangulare uniangulare, Tornoceras . . . 1, 10-12, 79, 158, 160, 163, 165,

254; Text-figs 23, 54A-C, 55A; Table 23; Pl. 26, Figs 1-6uniangulare widderi, Tornoceras . . . . . . . . . . . . . . 81, 254; Pl. 26, Fig. 5vagans, Carinoceras . . . . . . . . . . . . . . . 43, 48, 54-55, 79, 84-85, 138-140,

249; Text-figs 43A-E, 44D, 46A; Table 16; Pl. 23, Figs 5-16vagans, Carinoceras aff. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . 55, 79, 138, 141, 249; Text-fig. 23; Pl. 23, Figs 1-4vericosum, Archoceras . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 85Virginoceras Ruzhencev, 1960 . . . . . . . . . . . . . . . . 73, 122; Text-fig. 21Virginoceratinae Yatskov, 1990 . . . . . . . . . . . . . . . . . . . . . . . . . . . . 122wabashense, Archoceras . . . . . . . . . . . . . . . . . . . . . . . . 237; Pl. 17, Fig. 1Wedekindella Schindewolf, 1928 . . . . . . . . . 73, 76, 162; Text-fig. 21Wellsites House & Kirchgasser, 1993 . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . 49, 73, 77, 79, 84, 92, 95-96; Text-fig. 21


Abbey Gulf Beds (Penn Yan Shale) . . . . . . . . . . . . . . . . . 16, 20Angola Shale [marker (Scraggy Bed) black shale at base; top Rhinestreet

Shale] . . . . . . . . . . . . . 48-49, 54-55, 57; Text-figs 3-4, 13, 15Angola Shale (microcycles/minor rhythms abca; 1-9 of lower part) . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . 51-52, 65, 85; Text-figs 15, 19Angola Shale (West Falls Group) . . . . 4-5, 6, 43, 45, 48-49, 51-52, 54-

55, 64-65, 68-69, 79-80, 84-85, 123, 134-144, 148, 157, 166-169, 175-180, 184, 234, 238, 242, 245-246, 249, 265-266, 269;Text-figs 2-3, 4, 15, 23-24, 41-47, 53, 60, 63; Pl. 16, Figs 1-10,Pl. 18, Figs 3-4, 6-9, Pl. 20, Figs 1-2, 9-12, Pl. 21, Figs 1-12,Pl. 22, Figs 1-8, Pl. 23, Figs 1-4, 7-12, 15-16, Pl. 31, Figs 2-3,5-6, Pl. 32, Figs 2-14, Pl. 33, Figs 1-4, 9-20

Angola Shale equivalents . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . 48-49, 51, 57, 68; Text-figs 2-3, 4, 15, 41

Apulia Member (Lower Tully Formation) . . . . . . . . . . . . 11; Text-fig. 6Arkona Shale (Ontario, Canada) . . . . . . . . . . . . . . . . . . . . . . . . . . . 81Assise de Frasnes, lower (Europe) . . . . . . . . . . . . . . . . . . . . . . . . . . 70Assise de Matagne (Belgium) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 168Beards Creek Horizon (BC; West River Shale) . . . . . . . . . . . . . . . . . . . .

31-33, 35, 79, 102, 114-117, 122; Text-figs 9, 23, 33-34Bed G, West Brook Member (Upper Tully Formation) . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10-11; Text-fig. 6Bedford Shale (Conewango Group) . . . . . . . . . . . . . . . . . . . . . . 62, 87Beers Hill Member (Rhinestreet Shale) . . . . . . . . . . . . . . . . . . . . . 50-51Belgian F2d reefs (Belgium) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70Belgian F2h reefs (Belgium) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70Bellona Coral Bed (Tully Formation) . . . . 10-11, 66, 82; Text-figs 5-6Bergen Beach Bed (Lower Hubbard Quarry Submember Geneseo Shale)

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23Bituminous (Black) shale (of Clarke, 1904; Rhinestreet Shale) . . . . . . . 43Black Shale and Slate (Genesee Group) . . . . . . . . . . . . . . . . . . . . . . . 12Bluestone Bed (Ithaca Shale and Sandstone) (Genesee Group) . . . . 17, 26Bluff Point Silstone (BP; West River Shale) . . . . . . 30-34, 65, 79, 83,

103, 114, 117, 121, 218; Text-figs 23, 33-34; Pl. 8, Figs 10-11,1 4

Brallier Shale (or equivalents; Pennsylvania, West Virginia, Virginia) . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 157

Büdesheimer Schiefer (Germany) . . . . . . . . . . . . . . . . . . . . . . . . 167-168,172, 217, 257; Text-fig. 58; Pl. 7, Figs 2-3, Pl. 27, Figs 13-14

Burlington Limestone (Missouri) . . . 94; Text-fig. 27; Pl. 4, Figs 1-4Callaway Limestone (Missouri) . . . . . . . . . . . . . . . . . . . . . . . . . . . . 94Canadaway Group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . 4, 9, 57, 59, 161-163, 173, 179-180, 182; Text-fig. 2Cashaqua Shale (marker black shales A-F) . . 35-37, 38-41; Text-fig. 10Cashaqua Shale (Sonyea Group) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . 1, 3-5, 6, 34-35, 36-37, 38-42, 48-50, 64, 67-68, 79-80, 83, 93-95, 103-106, 108-110, 115, 125-127, 129-132, 152-154, 157, 165, 177, 184, 186, 206, 214, 217, 221-222, 225-226,229-230, 233-234, 250, 253, 265; Text-figs 2-3, 4, 10, 19, 23-24, 27H, 30E-H, 31A-H, 35, 36A-G, 37A-H, 38, 39A-J, 40,50A-Q, 51A-E, 54G, 55, 63; Pl. 2, Figs 1-2, 4-6, Pl. 6, Figs 1-

15, Pl. 9, Figs 2-3, 6-8, Pl. 10, Figs 1-4, 10, 12, Pl. 12, Figs 1-7, 11, Pl. 13, Figs 1-15, Pl. 14, Figs 1-10, Pl. 16, Fig. 12, Pl. 24,Figs 6, 8-9, Pl. 25, Fig. 1, Pl. 31, Figs 7-17

Cashaqua Shale (upper part or division of Kirchgasser, 1975) . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . 36-37, 68, 105-106, 109-110,157, 222, 225-226, 233-234, 258, 265; Text-figs 10, 30E-H,31A-H, 32, 38, 54-55; Pl. 6, Figs 1-15, Pl. 7, Figs 4-6, Pl. 10,Figs 5-9, 11, Pl. 11, Figs 1-13, Pl. 12, Figs 8-10, Pl. 15, Figs 1-2, Pl. 16, Fig. 11, Pl. 28, Figs 6-10, Pl. 31, Figs 18-20

Cashaqua Shale/Rhinestreet Shale contact (Sonyea/West Falls Groups) . .. . . . . . . . . . . . . . . . . . . . . . . . . 5, 34, 36-37; Text-figs. 3-4, 10

Catskill (Delta, clastic sequences; facies; red-bed succession; marine/non-marine) . . . . . . . . . . . . . . . . . . . . . 1-3, 63-64, 84; Text-fig. 18

Cattaraugus Shale and Sandstone (Conewango Group) . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . 4, 80, 87, 183-184; Text-figs 2, 24, 63

Cayuta Shale, Upper (Chesney Formation; Member, Chemung Group/Formation; Rhinestreet Shale equivalent) . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . 50, 96-98, 206, 209; Pl. 2, Fig. 8, Pl. 3, Fig. 1

Centerfield Limestone (Hamilton Group) . . . . . . . . . . . . . . . . . . 81-82Chadakoin Shale (Ellicott Shale, Coneaut Group) . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . 4, 40, 80, 86, 183-184; Text-figs 2, 24, 63Chagrin Member (Conneaut Group) . . . . . . . . . . . . . . . . . . . . . . . . . 62Chemung Group (Formation; rocks) . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . 5, 26, 51, 98, 107, 140; Text-figs 3-4; Pl. 2, Fig. 3Chemung Group (Grimes Sandstone at base; Naples, Ontario County) .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5, 140; Text-fig. 3Chemung Group, Lower . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26, 98, 107Cherry Valley Limestone (Marcellus Formtion, Hamilton Group) . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 81, 161-162Chittenango Shale (Marcellus Formation, Hamilton Group) . . . 81, 163Cleveland Shale (Ohio; Conewango Group) . . . 62, 87, 183; Text-fig. 61FConewango Group . . . . . . . . . . . . . . . . 3-4, 9, 62, 183; Text-figs 2, 61EConneaut Group (Formation) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . 4, 9, 62, 183, 273; Text-fig. 2; Pl. 35, Figs 11-13Conodont Bed (of Hinde, 1879; = North Evans Ls; Genesee Group) . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13, 27Corning Shale (Member, Rhinestreet Shale) . . . . . . . . 4, 49; Text-fig. 2Corell’s Point Goniatite Bed (CP; Gowanda Shale) . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . 5, 60, 79, 158, 174-175, 182, 257; Text-figs 3, 17, 23, 54M-O, 59B-C, 60A-C, 61A-B; Pl. 27, Figs 7-10, Pl. 29, Figs 1-15, Pl. 30, Figs 2, 4-14, Pl. 35, Figs 1-10

Crosby Sandstone (CS; Penn Yan Shale) . . . . . . . . 12, 17, 21-25, 28, 30,65, 79, 82, 103, 107, 113, 115, 117, 119, 165-166, 218, 254;Text-figs 7, 23, 34F, J, 54L; Pl. 8, Figs 6, 9, Pl. 26, Figs 7, 10

Dasberg (unit in Upper Devonian of Germany) . . . . . . . . . . . . . . . . . 77Domanik Suite, Lower (Group; Timan, Russia) . . . . . . . . . . 70, 108-109Dunkirk Shale (Black Shale; Canadaway Group) . . 1, 4, 5, 57, 59, 63-64,

68, 70-71, 79-80, 86,149, 184; Text-figs 2-3, 16, 23-24, 63Dunn Hill (Member, Rhinestreet Shale) . . . . . . . . 4, 49, 51; Text-fig. 2Ellicott Shale (Chadakoin Shale, Coneaut Group) . . . . . . . . 4, 62, 79, 86,

182-183, 273; Text-figs 2, 23, 61C-D, 62; Pl. 35, Figs 11-13

INDEX OF STRATIGRAPHIC UNITS

Pages denoting principal discussions, illustrations of units in text-figures, and illustrations of goniatites from units (text-figures and plates) are in boldfont.

W. tynaniHouse & Kirchgasser, 1993 . . . . . . . 50-51, 79, 83-84,94, 96, 185, 205; Text-figs 23, 27A, 63; Pl. 1, Fig. 9

W. williamsi (Wells, 1956) [Beloceras] . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . 50, 79, 94-96; Text-figs 23, 27B

WernerocerasWedekind, 1918 . . . . . . . . . . . . . . . . . . . . . . . . . . . . 76, 162widderi, Tornoceras uniangulare . . . . . . . . . . . . . . 81, 254; Pl. 26, Fig. 5williamsi,Wellsites . . . . . . . . . . . . . . . 50, 79, 94, 96; Text-figs 23, 27B


Enfield Shale (Formation, Sonyea Group) . . . . . . . . . . . . . . . . . . . . . . . .. . . 4-5, 43, 50, 65, 95, 109; Text-figs 2-3; Pl. 3, Figs 2-3, 5-6

Farnham Creek Bed (turbidite unit, lower Angola Shale) . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . 51-52, 54-55, 65; Text-fig. 15

Filmore Glen Bed(s) (Member, Tully Limestone) . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . 9-10, 11, 82, 92; Text-figs 5-6

Fir Tree Limestone (Pyrite; Submember, Geneseo Shale) . . . . . . . . . . . . .. . . . . . . . . . . . . . . 12-13, 16, 23, 27-28, 102, 186; Text-fig. 7

Firestone Bed(s) (facies; of Williams, 1884; Ithaca Shale and Sandstone) .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26, 108

Fossil Log Horizon (FL; Williamsburgh Bed; West River Shale) . . . . 31-33, 67, 79, 102-105, 122, 136; Text-figs 9, 23, 30D, 33K-L

Fossil Log Horizons . . . . . . . . . . . . 38, 40, 67, 79, 110; Text-fig. 23Gardeau Shale (and Sandstone; Rhinestreet equivalent in part; West Falls

Group) . . 4-5, 45, 48, 50, 65, 84, 148; Text-figs 2-3, 4, 12, 14Genesee Group (Formation) . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1, 4-5,

7, 12-13, 14-15, 17, 19-20, 33-35, 64-66, 69, 79, 99, 102-103,105-109, 112, 114-116, 119, 124, 130, 136, 158, 163-164, 166,170-172, 186, 221; Text-figs 2-3, 4, 7-9, 14, 23, 28A-K, 32,33A-O, 34, 54A-C, E-F, 57A, D-F; Pl. 9, Figs 1, 4-5

Genesee Slate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12, 34Geneseo Limestone Horizon (Geneseo Shale) . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . 1, 13-15, 16, 18-20, 102; Text-fig. 8Geneseo Shale (black shale; Member, Genesee Group) . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . 1, 4-5, 6, 9-10, 12-15, 16, 18-21, 23-24, 27, 63-66, 79-80, 82, 92, 99, 102, 164, 169-171,184, 186, 210, 265, 270;Text-figs 2-3, 4-7, 8, 23-24, 54E, 57D-F, 63; Pl. 4, Fig. 6, Pl. 31, Figs 14-15, Pl. 34, Figs 6-7

Geneseo (Shale)-Penn Yan (Shale) contact . . . . . . . . . . . . . . 5, 12-16, 18-21, 23-24, 79-80, 102, 171, 184; Text-figs 2-3, 7-8, 23-24, 63

Genundewa Limestone (Genesee Group) . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . 1, 4-5, 12-13, 14-15, 17-22, 27-31, 66-68, 70, 79-80, 82-83, 91, 103-107, 112-113, 115, 117, 119-121, 123-124,130, 132, 135-136, 157-158, 162-165, 172, 184, 186, 218, 237-238, 241, 250, 258; Text-figs 2-3, 4, 7-9, 23-24, 30A-C, I-L,33F-G, I-J, N-O, 34G, 35, 42F, 53A-B, 54F, 55, 63; Pl. 8, Figs4-5, 7-8, 12-13, Pl. 17, Figs 5-10, Pl. 18, Figs 1-2, 5, 11-12, Pl.19, Figs 10-16, Pl. 24, Figs 1-4, Pl. 28, Figs 1-5

Genundewa Limestone (Lower division) . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . 13, 14-15, 17, 27-30, 33, 82, 106,112-113, 115, 117, 119-121, 123-124, 164, 218; Text-figs 8-9

Genundewa Limestone (Upper division) . . . . . . . . . . . . . . . . . . . . . . . . .. 13-14, 15, 17-18, 27-30, 33, 83, 123-124, 136; Text-figs 8-9

Genundewa Limestone disconformity (sub-Upper Division) . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . 12-15, 18, 27; Text-figs 7-8

Genundewa Pyrite (Horizon) (pyritic nodular bed; Geundewa Limestoneequivalent) . . . . . . . . . . 12, 30, 105, 107, 120-121; Text-fig. 7

Goniatite Concretionary Layer (Parrish Limestone, Cashaqua Shale) . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42

Gogo Formation (Western Australia) . . . . . . . . . . . . . . . . . . . . . 109, 157Gorge Gully Submember (Windom Shale) . . . . . . . . . . . . . . . . . . . . . 11Gowanda Shale (Canadaway Group) . . . . . . . . . . . 3-5, 6, 59-60, 61-62,

68, 71, 80, 86, 158, 161-163, 173, 185-176, 179-182, 184, 261-262; Text-figs 2-3, 4, 17, 24, 54M-O, 59A-C, 60A-C, 61A-B,63; Pl. 29, Figs 1-15, Pl. 30, Figs 1-15, Pl. 31, Fig. 1

Grimes sandstone member (of Chemung formation of G. Chadwick) . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 140

Grimes Siltstone (Rhinestreet equivalent; West Falls Group) . . . . . . . . . .. . . . . . . . . . . 5-6, 45, 140, 249; Text-fig. 3; Pl. 23, Figs 13-14

Hamilton Group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . 4-5, 12-13, 17, 158, 161-164, 169; Text-figs 2-3, 54D

Hanover Shale (West Falls Group) . . . . . . . . . . 3-5, 6, 56-57, 58-59, 67,70, 79-80, 85-86, 134, 141, 149-152, 155, 175-176, 178-180,

184, 241, 265; Text-figs 2-3, 4, 16, 23-24, 41G, 48C, G, 49A-E, 60E, 63; Pl. 19, Figs 1-9, Pl. 31, Fig. 3, Pl. 32, Fig. 1

Hanover Shale/Dunkirk Shale contact (West Falls Group/CanadawayGroup) . . . . . . . . . . . . . . 5-6, 56, 58-59, 86; Text-figs. 3-4, 16

Hanover Shale equivalents (West Falls Group) . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . 5-6, 58-59; Text-figs 3-4, 16

Harrell Shale (Pennsylvania, West Virginia) . . . . . . . . . . . . . . . . . . . . . . .. . . 68, 106, 112-113, 117, 120, 170-172; Text-figs 57B-C, 58E

Hatch Shale (Rhinestreet equivalent, West Falls Group) . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . 5-6, 44-45; Text-figs 3-4, 11

Hemberg (unit in Upper Devonian of Germany) . . . . . . . . . . . . . . . . . 77Hiatus concretion horizon (of Baird, 1976; Penn Yan Shale) . . 17, 21, 65High Point Sandstone (= Nunda Sandstone at Naples, Ontario County).

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 140High Point Sandstone (West Falls Group) . . . . . . . . . . . . . . . . . . 95, 140Hubbard Quarry Shale (Submember, Geneseo Shale) . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . 13, 16, 23-24, 169, 171; Text-fig. 57EHuddle Rider Bed (black shale in upper Hanover Shale) . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . 56, 59, 71, 86; Text-fig. 16Hungry Hollow Formation (Ontario, Canada) . . . . . . . . . . . . . . . 81-82Ithaca Beds (of Chemung Group) . . . . . . . . . . . . . . . . . . . . . . . . . . . . 107Ithaca Shale and Sandstone (Formation, Genesee Group) . . . . . . . . . . . .

. . . . . . . . . . 4-5, 6, 12, 16-17, 21-24, 26-28, 30, 34, 65, 82-83,100, 107-109, 113, 115, 119, 121, 170-171, 186; Text-figs 2-3,4, 28H-K, 33I-J, 57A, C; Pl. 9, Figs 1, 4-5, Pl. 34, Fig. 8

Java Formation (West Falls Group) . . . . . . . . . . . . . . . 4, 57; Text-fig. 2Kellwasserkalk (Lower, Upper; Germany) . . . . . . . . . . . . . . 70-71, 85-86Knollenkalk facies . . . . . . . . . . . . . . 5, 35, 55, 57, 59, 66-67; Text-fig. 3Kramenzelkalk facies . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66-67Laona Sandstone (Siltstone; Canadaway Group) . . . . . . . . . . . . . . . . . . .

. . . . 4-5, 6, 59, 61, 86, 180; Text-figs 2-3, 4; Pl. 30, Figs 1-3Leicester Pyrite (base of Geneseo Shale, Genesee Group) . . . . . . . . . 3-5,

9-10, 12-13, 14-15, 16-20, 69, 81-82, 158-160, 163-164, 186,254; Text-figs 2-3, 4, 7-8, 54A-C, 55A; Pl. 26, Figs 1-4, 6

Linden Horizon (LH; Linden Goniatite Horizon; Penn Yan Shale) . . . . .. . . . . . . . . 14-15, 17-18, 19-23, 26, 79, 82, 103, 107, 115-116,119, 164, 186; Text-figs 8, 23, 34A, C-D; Pl. 8, Figs 3-4

Lodi Limestone (LO) . . . . . . . . 5-6, 12-13, 14-16, 18-21, 23, 24-25, 28,66, 79-80, 82, 99-102, 164, 184, 186; Text-figs 3-4, 7-8, 23-24, 28B-E, G, 29A-E, 32, 63; Pl. 4, Figs 5, 7, Pl. 5, Figs 1-9

Lodi Limestone beds A-D . . 16, 23-24, 28, 101-102; Text-fig. 29A-ELower Black Band (Middlesex Shale) . . . . . . . . . . . . . . . . . . . . . . 34, 95Lyaiol Formation (Timan, Russia) . . . . . . . . . . . . . . . . . . . . . . . . . . . 176Marcellus Formation (Hamilton Group) . . . . . . . . . . . . . . . . . . 162-163Middlesex Shale (Black Shale; Sonyea Group) . . . . . 1, 4-5, 6, 12, 17, 30-

34, 35-36, 37, 40, 42, 49-50, 63-64, 67, 69-70, 79-80, 83-84,92-93, 95, 102, 105, 115, 117, 121-122, 136, 184, 206; Text-figs 2-3, 4, 9-10, 23-24, 63; Pl. 2, Fig. 7, Pl. 3, Figs 5-6

Middlesex Shale equivalents (Sonyea Group) . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . 33-34, 49-50, 83, 93, 95, 206; Pl. 2, Fig. 7

Millboro Shale (Virginia) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 157Millport Member (Rhinestreet Shale) . . . . . . . . . . . . . . . . . . . . . . . . . 51Montour Shale [Lower tongue (Member) of Middlesex Shale] . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4, 34, 49; Text-fig. 2Moravia Bed (Bed G, West Brook Member, Tully Formation) . . . . . . . . .

. . . . . . . . 9-11, 66, 82, 91, 205; Text-figs 5-6; Pl. 1, Figs 1-8Moreland Shale (Lower Member, Rhinestreet Shale) . . . . . . . . . . . . . . . .

. . . . . 4, 49-51, 84, 96, 98, 205; Text-figs 2, 27A; Pl. 1, Fig. 9Moscow Shale (Formation, Hamilton Group) . . . . . . 4-5, 10-11, 12-13,

17-20, 79-80, 164, 184, 284; Text-figs 2-3, 6-7, 8, 23-24, 63Moscow-Geneseo (Hamilton/Genesee) contact . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . 4-5, 10, 12, 14-15, 18-21; Text-figs 2-3, 4, 7-8Naples Group, Beds (beds, facies, Formation) . . . . . . . . . . . . . . . . . . . . .


. . . . . 5, 34, 64, 83, 95, 110, 115, 123, 125-127, 132, 158, 161,166, 218, 221, 225, 237, 246; Text-figs 3, 31A, 34A, 36A-C,54G; Pl. 8, Figs 1-2, Pl. 9, Figs 2-3, Pl. 11, Figs 6, 12-13, Pl. 14,Fig. 3, Pl. 17, Fig. 5, Pl. 22, Figs 4-5, Pl. 27, Fig. 15

Nehden (Nehdener Schichten/Schiefer; -Schurbusch; Germany) . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3, 71, 77, 181

New Albany Shale (Indiana) . . . . . . . 237; Pl. 17, Fig. 1, Pl. 18, Fig. 10New Albany Shale (Trousdale Member, Kentucky) . . . . . . . . . . . . . . . 186North Evans Limestone (Conodont Bed of Hinde, 1879; Genesee Group)

. . . . . . . . . . . . . . . . . 12-15, 17-18, 27-28, 64; Text-figs 7-8Nunda Sandstone (and equivalents; West Falls Group) . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4-5, 6, 45, 51-53, 55-57,59, 68, 108, 140, 143, 145, 148; Text-figs 2-3, 4, 15, 45C-D

Old Red Sandstone (England; Europe) . . . . . . . . . . . . . . . . . . . . . . 3Oswayo Shale and Sandstone (Conewango Group) . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . 4, 80, 183-185; Text-figs 2, 24, 61E, 63Panama Conglomerate (Conewango Group) . . . . . . . . . . . . . . . . . . 4, 62Parrish Limestone (PL; Cashaqua Shale) . . . . . . . . . . . . . . . . . . . . . . . . .

. . . 5-6, 35-37, 41-42, 43, 68, 79, 125-127, 129-130, 152-153,155, 157, 222; Text-figs 3-4, 10, 23, 36D-G, 37D, G-H, 50H,K, M-N, P, 51E; Pl. 10, Figs 1-2, 12, Pl. 24, Figs 5-9

Penn Yan Black Shale A . . . . . . . . . . . 14-15, 18-19, 20, 40; Text-fig. 8Penn Yan Black Shale B . . . . . . . . . . . . . . . . 14-15, 18-19; Text-fig. 8Penn Yan Shale (Genesee Group) . . . . . . . . . . . . . . . . . . . . . . . . . . . 4-5,

6, 12-13, 14-15, 16-24, 26-29, 33, 65, 68, 79-80, 82, 100-103,107-109, 112-113, 115, 117, 119-120, 123, 164, 166, 170-171,184, 218, 270; Text-figs 2-3, 4, 7-9, 23-24, 28F, 34A-F, J, 35A,57D, 63; Pl. 8, Figs 1-4, 6, 9, Pl. 34, Figs 6-7

Penn Yan (Shale)-Crosby (Sandstone) contact . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . 12, 17, 22-23, 65; Text-fig. 7

Penn Yan (Shale)-Ithaca (Shale and Sandstone) contact . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . 5-6, 12, 22-23, 65; Text-figs 3-4, 7

Pentonwarra Goniatite Bed (North Cornwall, England) . . . . . . . . . . 76Perrysburg Formation (Canadaway Group) . . . . . . . . . . 4-5; Text-fig. 3Pharciceras Bed (West Brook Member, Tully Limestone) . . . . . . . . . . . . .

. . . . . . . . . 10-11, 158, 205; Text-figs 6, 54K; Pl. 1, Figs 4, 7Pharciceras horizons (Trousdale Member, New Albany Shale, Kentucky) .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 186Pharciceras-Lage [Untere; Martenberg section, Diemelsee (Adorf ), Ger-

many] . . . . . . . . . . . . . . . . . . . . . . . . . . . . 91; Text-fig. 26A-CPipe Creek Shale (Black Shale) (Formation,West Falls Group) . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1, 4-5, 6, 49, 55-57, 58-59, 63, 67, 70, 79-80, 85, 184; Text-figs 2-3, 4, 16, 23-24, 63

Pipe Creek Shale/Hanover Shale contact . . . . . . . . 5, 56; Text-figs 3, 16Platyceras Bed (West Brook Member, Tully Limestone) . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 158, 165; Text-fig. 54KPoint Breeze Goniatite Bed (PB; Angola Shale) . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . 52-54, 55-57, 79, 84-86, 131,134-144, 148, 166-169, 175-176, 179; Text-figs 15, 23, 41-44,47, 53, 60; Pl. 16, Figs 3-8, Pl. 20, Figs 1-6, 9-10, Pl. 23, Figs7-12, Pl. 32, Figs 4-9, 11-14, Pl. 33, Figs 3-4, 9-14, 19-20

Portage (Ithaca beds) at the University Quarry (Ithaca, Tompkins County). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 108; Pl. 9, Fig. 1

Portage (Naples) Beds . . . . . . . . . . . . . . 115, 225; Pl. 11, Figs 6, 12-13Portage (rocks, beds, shales, Group; outcrops in town of Naples; sandstone

on East Hill, Naples, Ontario County; Java, Wyoming County;Eighteenmile Creek, Erie County) . . . . . . . . . . 3, 34, 62, 95, 123,131, 139, 153, 166; Text-figs 43-44, 50; Pl. 23, Figs 13-14

Portage Group (Ithaca, Tompkins County) . . . . . . 108; Pl. 34, Fig. 5Portage Group (Lower) (Genesee River) . . . . . . . . . . . . . . . . . . . . . 45, 57Portage Sandstone (Genesee River) (West Falls Group) . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 140, 148; Text-fig. 44Prout Limestone (Erie County, Ohio) . . . . . . . . . . . . . . . . . . . . . . . . . 81

Pulteney/Rock Stream turbidite wedge (Cashaqua Shale equivalents) . . .. . . . . . . . . . . . . . . . . . . . . . . 4-5, 6, 34-37; Text-figs 2-3, 4, 10

Pulteney Shale (Member, Cashaqua Shale) . . . . 4-5, 6, 34; Text-figs 3-4Quarry Sandstone interval (Ithaca Formation) . . . . . . . . . . . . 17, 26, 109Relyea Creek Horizon (RL) (upper Rhinestreet Shale). . . . . . . . . . . . . . .

. . . . . . . . . . . . 43, 46-49, 79, 84, 134, 140-141, 143-144, 148,166-167; Text-figs 13, 23, 41F, 44F, 45B; Pl. 23, Figs 5-6

Renwick Shale (RE; Black Shale; Formation, Genesee Group) . . . 1, 4-5,6, 12, 16-18, 20-26, 79, 82, 102, 108, 186; Text-figs 2-3, 4, 23

Rhinestreet Shale (Black Shale; West Falls Group) . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . 4-5, 6, 34-35, 38, 40, 43-47, 48-50,55, 57, 64, 66-67, 69-70, 79-80, 83-84, 94-98, 110, 134-136,138-140, 142-145, 148, 156, 161, 166-167, 176, 184, 186, 205-206, 209, 242, 246, 249, 253, 257, 265, 269; Text-figs 2-3, 4,10-11, 13, 15, 23-24, 27A-B, I, 41B, F, 42A-B, E, 43D-E, 44F,45B, 46-47, 52A-H, 56A-D, 63; Pl. 1, Fig. 9, Pl. 3, Fig. 1, Pl.20, Figs 7-8, Pl. 22, Fig. 9, Pl. 23, Figs 5-6, 13-14, Pl. 25, Figs2-5, Pl. 27, Fig. 12, Pl. 31, Fig. 4, Pl. 33, Figs 5-8

Rhinestreet Shale (microcyles of upper part) . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . 43-44, 46-47, 69; Text-fig.13

Rhinestreet Shale/Angola Shale contact . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . 5-6, 46-47, 52-53; Text-figs 3-4, 13, 15

Rhinestreet Shale equivalents (West Falls Group) . . . . . . . . . . . . . . . . . .. 5-6, 44-45, 46-47, 49-40, 57, 84, 97-98; Text-figs 3-4, 11-13

Rock Stream Siltstone (Sonyea Group) . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . 4-5, 6, 34-37, 40-43, 68; Text-figs 2-3, 4, 10

Rock Stream Silstone/Cashaqua Shale contact . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . 5, 36-37, 41-42; Text-figs 3, 10

Roricks Glen Shale (Member, Rhinestreet Shale) . . . . . . . . . 4; Text-fig. 2Rye Point Shale (Member, Cashaqua Shale) . . . . . . . . . 4, 35; Text-fig. 2Sawmill Creek Shale (Upper Member, Middlesex Shale) . . . . . 34, 49-50Saltern Cove Goniatite Bed (Devon, England) . . . 167; Text-fig. 56E-FSB (Schumacher) black shale (Penn Yan Shale) . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . 14-16, 17-21, 102; Text-fig. 8Schistes de Matagne (Matagne Shale; F3; Belgium) . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . 70, 140, 149-150, 168; Text-fig. 44B, 48BSchumacher Bed (SB; concretion bed, Penn Yan Shale) . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14-16, 18; Text-fig. 8Scraggy Bed (concretion bed; marker black shale; top of Rhinestreet Shale)

. . . . . . . . 5-6, 43-44, 46-47, 48, 54, 148; Text-figs 3-4, 13, 15Second black band (of Clarke, 1898; Rhinestreet Shale) . . . . . . . . . . . 43Sherburne Siltstone (Formation, Genesee Group) . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . 4-5, 6, 12, 16, 21, 24-27, 99-100, 102,210; Text-figs 2-3, 4, 7, 28A; Pl. 4, Figs 7-9, Pl. 5, Figs 1-9

Shurtleff Septarian Horizon (SH; Upper division, Cashaqua Shale) . . . . .. . 5-6, 35-38, 39-40, 41-42, 67-68, 79, 83, 103-105, 110, 125,127, 129-130, 157, 165, 177, 214, 217, 222, 225-226, 233-234,258, 265; Text-figs 3-4, 10, 23, 30E-H, 31A, C-E, G-H, 32B,38C, 55C; Pl. 6, Figs 1-15, Pl. 7, Figs 4-6, Pl. 10, Figs 5-9, 11,Pl. 11, Figs 1-13, Pl. 12, Figs 8-10, Pl. 15, Figs 1-2, Pl. 16, Fig.11, Pl. 28, Figs 6-10, Pl. 31, Figs 18-20

Sonyea Group . . . . . . . . . . . . . . . . . . 4-5, 6, 8, 17, 34-35, 36-37, 49-50,63-64, 68-69, 79, 92-93, 95, 103-105, 108-110, 115, 129, 130,132, 137, 157-158, 165, 177; Text-figs 2, 3-4, 10, 14, 18, 23

Sonyea Group (major rhythm ABCBA) . . . . . . . . . . . 69; Text-fig. 19ASonyea Group equivalents (south and southeast of Ithaca, Tompkins

County) . . . . . . . . . . . . . . . . . . . . . . . . 43, 49-50; Text-fig. 14South Wales Shale (Canadaway Group) . . . . . . . . . . . . . . . . . . . . . . . . 59Squaw Bay Limestone (Michigan) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . 68, 112-113, 117, 120, 158, 165-166,170, 254, 257; Text-fig. 54H-J; Pl. 26, Figs 8-9, Pl. 27, Fig. 16

Starkey black shale (Penn Yan Shale) . . . . . . . . . . . . . . . . . . . . . . . . . . 21Styliola Limestone (Genundewa Limestone) . . . . . . . . . . . . . . . 27, 115


Table Rock Sandstone (Genesee River) (Gardeau Shale,West Falls Group). . . . . . . . . . . . . . . . . . . . . . . 5-6, 45, 57, 148; Text-figs 3-4, 12

Taghanic Onlap (Sequence) . . . 5, 9, 12, 63, 66, 70, 186; Text-figs 3, 7Taghanic Unconformity . . . . . . . . . . . . . . . . . . . . 9, 12-13; Text-fig. 7Trinity Horizon (TR; “The Trinity” black shales; Angola Shale) . . . . . . .

. . . . . . . . . . . . . . . . 51-53, 55, 57, 68, 79, 85; Text-figs 15, 23Tully Limestone (Formation) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . 1-5, 6-7, 9-10, 11-12, 16, 24, 27, 63, 66, 70, 78-79, 80-82, 89-92, 158, 165, 170, 184, 186, 205; Text-figs 2-3, 4-5, 6-7, 23-24, 26D-H, 54K, P, 63; Pl. 1, Figs 1-8, Pl. 27, Fig. 11

Tully Limestone (Upper Member/Division) . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . 9-10, 11-12, 27, 66, 70, 82,186, 205, 257; Text-figs 5-7; Pl. 1, Figs 1-8, Pl. 27, Fig. 11

Tully Limestone disconformities (pre-intra-post Tully disconformities) . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . 9-10, 12, 78; Text-figs 5-7

Tully Limestone-Geneseo Shale contact . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . 5-6, 9-10, 12; Text-figs 3-4, 5-7

Tully Pyrite (Leicester Pyrite) . . . . . . . . . . . . . . . . . . . . . . . 158-159, 163Union Spring Shale (Werneroceras Bed; Marcellus Formation, Hamilton

Group) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 162Upper black shale (of Hall, 1843, of Reports; Rhinestreet Shale) . . 12, 43Venango Sandstone, Lower (Cattaraugus; Conewango Group; Erie

County, Pennsylvania) . . . . . . . . . . . . . . . 4, 62, 87; Text-fig. 2Virgin Hills Formation (Western Australia) . . . . . . . . . . . . . . . . . 150, 155Volusia Member (Conneaut Group) . . . . . . . . . . . . . . . . . . . . . . . . . . 62Wanakah Shale (Member, Ludlowville Formation, Hamilton Group) . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 169Werneroceras Bed (Union Springs Shale, Marcellus Formation, Hamilton

Group) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 162West Brook Shale (Member, Bed, Tully Formation) . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . 9-10, 66, 70, 82, 91, 158, 165,205, 257; Text-figs 5-6, 54K, P; Pl. 1, Figs 1-8, Pl. 27, Fig. 11

West Falls Group (Formation) . . . . . . . . . . . . . . . . . . . . . . . . . 4-5, 6, 8,

34-35, 43, 51, 57, 65, 68-70, 79-80, 92, 95-98, 108, 123, 134-135, 137-142, 144-145, 148, 155, 161, 166, 178-179; Text-figs2-3, 4, 14, 23, 41A-G, 43A-g, 44A, C-F, 45B; Pl. 3, Fig. 4

West Falls Group (Formation), Lower (major rhythm ABCA) . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69; Text-fig. 19B

West Falls Group equivalents (south and southeast of Ithaca, TompkinsCounty) . . . . . . . . 49-50, 95, 97-98; Text-fig. 14; Pl. 3, Fig. 4

West Hill Member/Nunda Sandstone interval (West Falls Group) . . . . . .. . . . . . . . . . . . . . . . . . . . . . . 5-6, 45, 51, 55, 68; Text-figs 3-4

West Hill Shale (Flags; Sandstone; Member, Angola Shale) . . . . . . . . . . .. . . . . . . . . . . . . . . 5-6, 45, 51, 55, 68, 140, 148; Text-figs 3-4

West River Shale (23 cycles/rhythms) . . . 30, 33, 69-70; Text-fig. 9West River Shale (Formation, Genesee Group) . . . . . . . . . . . . 3-5, 6, 12,

27, 30, 31-33, 35, 65, 67-69, 79-80, 82-83, 102-107, 113-115,117, 121-122, 129, 136, 164, 172, 184, 217-218, 270; Text-figs2-3, 4, 7, 9, 23-24, 30D, 33A-E, H, K-M, 34H, K, 58A-B, 63;Pl. 7, Fig. 1, Pl. 8, Figs 10-11, 14, Pl. 34, Figs 1-4

Westfield Shale (Canadaway Group) . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . 4, 61, 180, 184, 262; Text-figs 2, 24; Pl. 30, Figs 1, 3

Widder Shale (Ontario, Canada) . . . . . . . . . . 81-82, 254; Pl. 26, Fig. 5Williams Brook Coquinite (Genesee Group) . . . . . 17, 24-25, 82, 108Williamsburgh Bed (Fossil Log Horizon, FL; West River Shale) . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31-32, 33,79, 102, 104-105, 122, 136; Text-figs 9, 23, 30D, 33K-L

Windom Shale (Member, Moscow Formation, Hamilton Group) . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9, 11-12, 17, 21, 158, 164, 257; Text-figs 7, 54D; Pl. 27, Figs 1-6

Wiscoy Sandstone (and equivalents) (West Falls Group) . . . . . . . . . . . . .. . . . . . . . 4-5, 6, 57-58, 59, 145, 148; Text-figs 2-3, 4, 45C-D

Wocklum (unit in Upper Devonian of Germany) . . . . . . . . . . . . . . . . 77Wolf Creek Conglomerate (Conewango Group) . . . . . 4, 62; Text-fig. 2Woodmont Shale (or equivalents; West Virginia, Maryland) . . . . . . . 157




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Documents

Late Devonian Goniatites (Cephalopoda, Ammonoidea) from New