Late Devonian Goniatites (Cephalopoda, Ammonoidea) from New York State

M. R. House and W. T. Kirchgasser

Bulletins of American Paleontology

Number 374, July 2008 BULLETINS OF AMERICAN PALEONTOLOGY Established 1895

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Th is paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). Late Devonian Goniatites (Cephalopoda, Ammonoidea) from New York State

M. R. House and W. T. Kirchgasser

Bulletins of American Paleontology

Number 374, July 2008 ISSN 007-5779 ISBN 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.

HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 1

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 York 13676, U. S. A., email [email protected] [*deceased]

ABSTRACT This 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; the taxonomy 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 of primary material at 184 localities. A description of the considerable additional goniatite material is integrated with that of the classic type specimens to treat the faunas systematically and as a whole. Noteworthy is the different contribution to knowledge provided by a range of preservation patterns, especially of the material preserved as barite replacements from the Cashaqua Shale. The goniatite-bearing horizons are mostly associated with transgressive pulses that punctuate the Catskill 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, and with the new international standard established by Becker & House (2000). Minor regional variations give a more detailed Appalachian regional zonation of 25 zones, which is integrated with the international standard. The succession associated with the major late Givetian faunal turnover (Taghanic Event) is described, and the influence of sea level changes and anoxia on faunal entry and diversification is reviewed. Very little endemicity is recognized in the faunas, apart from the multilobed Triainoceratidae that are well developed in the mid-Frasnian of central New York. The late Frasnian succession 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 beckeri n. 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 of Manticoceras 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. sinuosum tardum 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.

INTRODUCTION turbed, and over a wide outcrop tract more than 280 mi Nowhere in the world is the Upper Devonian exposed so (450 km) long, the dip of the rocks rarely exceeds 1.5o to continuously and accessibly as in New York State. The lower the south. With so extensive an outcrop, and so simple a Upper Devonian (upper Givetian-Frasnian) in particular is structure, the facies changes from the thin marine sequence seen in quite unparalleled excellence, and this interval has near Lake Erie on the west to the thick non-marine se- the greatest potential for refining the biostratigraphic record quences in the Catskill Mountains to the east can be corre- of goniatite cephalopods. The Upper Devonian in New York lated and analyzed in great detail (Text-fig. 1). State crops out over an area of approximately 15,000 mi2 These goniatite faunas have been known since the work (39,000 km2). The sequences are well exposed in innumer- of James Hall (1843) and Lardner Vanuxem (1839, 1842), able creek, gully, and riverside sections. The beds are undis- but it was their description by John M. Clarke (1898, 1899a, b) that made them particularly well known. Clarke 2 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

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 quadrangle maps, current editions of the New York State Atlas and Gazeteer (Anonymous, 1993) are a convenient source of maps for site location.

discovered very well-preserved material, including rich bar- The framework was built on the lithological correlations es- itized faunas that enabled extraction by etching, and he de- tablished by G. W. Colton, J. F. Pepper, W. de Witt, Jr., and veloped techniques for producing artificial molds of material R. G. Sutton in a series of contributions. So excellent are from siltstones. His descriptions of the early stages of many the exposures, however, that individual goniatite horizons forms, particularly species of Manticoceras, are still among have been traced, in some cases, for distances of more than the more interesting of goniatite ontogenetic studies. This 50 mi (80 km). Substantial detail has thus been added to early work was largely collated and revised by A. K. Miller the stratigraphical picture built up by many earlier geolo- (1938) in a review of American Devonian ammonoids, but gists. hardly any of Clarke’s material was reillustrated photograph- This work describes the occurrence and sequence of go- ically, and the original drawings were merely reproduced. niatite faunas for the lower Upper Devonian which is un- Modern illustration has long been needed. equalled in the world for its stratigraphical discrimination. The New York goniatite sequence has largely been ig- It contributes to a detailed documentation of goniatite evo- nored biostratigraphically, apart from broad reviews by lution during this period and adds detail on the facies House (1962, 1978). The early faunas described by Clarke changes associated with the goniatite horizons as they pass were from isolated localities, the relations of which have not eastward toward the Catskill Delta. been 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 a HISTORICAL REVIEW manner as to make their geographical position doubtful ex- When, in the autumn of 1836, James Hall and Lardner cept in the most general terms. Vanuxem commenced field work in the Devonian area for This study was commenced by the authors in 1964, pri- the newly created Geological Survey of New York, much had marily to relocate the old sources of material and to establish already been accomplished in the establishment of the their stratigraphic relations. This was an ambitious under- stratigraphy and distribution of the Devonian rocks, even taking and involved the study and measuring of many sec- though the System was not itself designated until 1839 tions in western upstate New York to document the facies (Sedgwick & Murchison, 1839a: 259; 1839b: 121). The pi- changes and discover the sequence of ammonoid faunas. oneer period of geological investigation, up to the establish- HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 3

ment of the Survey, has been reviewed in a thorough essay ditional goniatites, but with the arrival of J. M. Clarke at by J. W. Wells (1963). the Survey, substantial advances followed (Clarke 1882, With the publication of Part III of the Geology of New 1883, 1884, 1885). In 1885, Clarke described some mate- York (Vanuxem, 1842), and particularly of Part IV (Hall, rial from the Cashaqua Shale and discussed the goniatites 1843) with its geological map and systematic stratigraphical (Clarke 1891a, b). The supposed recognition of a clymeniid treatment, a firm foundation of geological knowledge was (Clarke, 1892) was mistaken, but the baritic preservation of established. Understandably, at that time, correlation with the material might well have led him to appreciate the value Europe was imprecise, and Hall thought that the Portage of acid-etching of New York material. Certainly it was the and Chemung rocks, now referred to the Upper Devonian, development of these techniques that resulted in his impor- preceded the Old Red Sandstone. Their Devonian age seems tant study of goniatites in The Naples Fauna (Clarke 1898, first to have been appreciated by Ferdinand Roemer in 1846 1899a, b). In this work much new material was described, (Clarke, 1921: 172; Wells, 1963: 64) who had visited Hall especially from the Cashaqua, Angola, and Hanover Shales, at Albany in 1845. but the locality details were poorly recorded; a useful source Hall had, however, recognized the broad pattern of of information on localities is a small guidebook by Clarke thickening from the marine rocks of the west to the terres- to fossiliferous localities in western New York (Clarke, trial facies in the east, and had recognized the extent of the 1899c). Had stratigraphical precision been given for the go- Tully Limestone, and of the major wedges of black shales, all niatite localities and faunas described by Clarke, there is lit- giving time horizons extending eastward to the thick Catskill tle doubt that he would have laid the foundation of Frasnian red-bed succession. Hall described Devonian goniatites from goniatite biostratigraphy. As he did not do this, that distinc- a horizon within the Tully Limestone up to faunas on Lake tion went to Denckmann, Frech, and especially Wedekind, Erie shore in Chautauqua County, now known to belong to working on the much less well-exposed rocks in the Rhenish equivalents of the German Nehdener Schiefer (early Fa- Schiefergebirge of Germany. mennian). The source of this material will be discussed in The early decades of the twentieth century saw many the next section. new records of goniatites, mostly in the bulletins accompa- Hall had the benefit of accompanying Charles Lyell over nying the geological quadrangle mapping of upstate New much of the ground covered by his survey before publication York. New forms, however, were not recognized. Loomis of The Geology of New York, Part IV (Friedman, 1998). Lyell (1903) described early stages of goniatites from the Leicester was the leading theoretical geologist of his time and we have Pyrite. Clarke (1902, 1903) added further notes, and the no way of assessing the contributions he made to the devel- second part of The Naples Fauna was published in 1904. opment of Hall’s ideas and vice versa. It is a matter of surprise The next significant discovery was of Paralegoceras? milleri that Hall did not consider with any accuracy the matter of (now Maeneceras milleri) by Flower & Caster (1935) in the correlation with Europe. By the time Hall met him, Lyell Conewango Group in neighboring Pennsylvania, although had already, as President of the Geological Society of Lon- it was not until a later paper (Miller & Flower, 1936), that don, presided over many of the bitter arguments related to the generic assignment to Sporadoceras was given and its the establishment of the Devonian System in Europe (Rud- recognition as an indicator of the mid-Famennian was ap- wick, 1985). There is no evidence in Hall’s 1843 volume preciated. that these problems had been resolved, nor of their relevance A useful compilation and revision of earlier work of New for the North American rocks. Only a correlation with the York Devonian goniatites was given by A. K. Miller in 1938. Old Red Sandstone was attempted, and that was related This took advantage of the substantial German work of the solely to the terrestrial Catskill facies. previous 30 years to give more up-to-date generic assign- Vanuxem (1842), in reporting on the third geological ments, but no fieldwork was done for this compilation, and district to the east of Hall’s fourth district, was already in most illustrations were merely copied from earlier sources, rocks of a more clastic facies, but he too described goniatites especially Clarke’s monographs of 1898 and 1899. from rocks that would now be termed Frasnian, or lower Subsequently, Koenenites was recognized in the West Upper Devonian. New names were given to other goniatites River Shale (Wells, 1956b), Cheiloceras in the Gowanda in the same year by Conrad (1842), one of the state geolo- Shale, and Pharciceras in the Tully Limestone (House, gists whom Hall replaced as State Paleontologist in 1843. 1962). The latter two genera are key markers in the Euro- So far as goniatites are concerned, investigations ceased pean biostratigraphy. Clarke’s supposed clymeniid was for a substantial time, apart from spasmodic records (Hall, shown to be a manticoceratid (House, 1961) and a broad 1874, 1875). During the course of his Paleontology of New correlation of North American and European goniatite York, Hall (1876, 1879; Hall & Clarke, 1888) described ad- zones was attempted (House, 1962, 1968, 1978). The 4 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

of the State. This led to some revisions of correlations. Nev- ertheless, the stratigraphic framework is built on the detailed work of G. W. Colton, W. de Witt, Jr., J. F. Pepper, and R. G. Sutton and the reviews of L. V. Rickard. The relation of our sequences to earlier work is discussed in the following text. Our locality numbers are shown in Text-fig. 4 and listed in Table 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 section descriptions, notably by de Witt and Colton. For locality references, we used the U. S. G. S. quadrangle maps; a useful modern guide to the localities is the New York State Atlas and Gazeteer (1993 and later editions). The terminology for the main lithostratigraphic units in western and central parts of New York State is shown in Text-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 of early beloceratids, here placed in the genus Naplesites, which was recorded from Naples (Ontario County) in a facies that Text-fig. 2. Lithostratigraphic divisions of late Devonian rocks of appears to be within the Rhinestreet Shale (see Addendum). New York State (based on Rickard, 1975), with chronostratigraphic A comment is required on measurements. The current boundaries of the stage divisions drawn following decisions of the In- topographic maps show only scale in miles and altitude in ternational Commission on Stratigraphy (ICS) and International Union of Geological Sciences (IUGS). feet. Thus we are obliged to give locations using English measurements, but metric equivalents are given in paren- theses. For measured sections, metric measurements are tornoceratid succession of New York formed the basis for an given, with English equivalents in parenthesis. For paleon- allometric study of the group (House, 1965). Substantial tological descriptions only metric measurements are given. new information on the biostratigraphic occurrences of Pro- A facies transect for the rocks with which we are con- beloceras and Ponticeras was given in a review of these genera cerned is shown in Text-fig. 3. An outline from this (Text- by Kirchgasser (1968, 1975), and details have been given of fig. 4) is used to indicate the most important of the sections goniatite levels in the Genesee Group (Kirchgasser, 1973, we have studied; those of earlier workers are referred to in 1982, 1985, 1994, 1996a, b, 1998) and Sonyea Group the text, as appropriate. Fuller details of localities, including (Cashaqua Shale) (Kirchgasser, 1965, 1967, 1969, 1975) relevant 7.5’ quadrangle maps and county location, are given and comparisons made with the conodont succession in Table 1. (Kirchgasser, 1974, 1994; Klapper et al., 1995). Finally, the present authors published general reviews of Frasnian faunas TULLY FORMATION of New York (Kirchgasser & House 1981; House & Kirch- The Tully Limestone is one of the main regional, base-line gasser, 1993) that contributed to the establishment of a new marker units for the interval examined here. The Tully is the international goniatite zonation for the Frasnian Stage last of the substantial calcareous intercalations that charac- (Becker et al., 1993, Becker & House, 2000). terize marine Devonian in New York State. The name was applied by Vanuxem (1839: 278; 1842: 285) for the se- STRATIGRAPHY quence near Tully (Onondaga County) and nearby areas. INTRODUCTION Near Tully it is over 9 m (30 ft)-thick (Trainer, 1932). East- This section reviews the levels at which late Devonian am- ward there is a passage into more clastic facies (Cooper & monoids have been discovered in New York State. To locate Williams, 1935; Cooper, 1968; Johnson & Friedman, 1969) the exact horizons of earlier collections, considerable effort and westward, although anomalously thick owing to a “reef” was expended in measuring sections across the western part or mound complex at Borodino, the thicknesses are reduced OS ICGSE:LT EOINGONIATITES DEVONIAN LATE KIRCHGASSER: & HOUSE

Text-fig. 3. Facies diagram indicating the main rock units in the late Devonian of New York State from which goniatites are described in this study. At top left, read Corell’s (for Correll’s) Point Goniatite Bed. 5 6 B LEISO MRCNPLOTLG o 374 no. PALEONTOLOGY AMERICAN OF ULLETINS

Text-fig. 4. Outline of the main rock units in the late Devonian of New York State and the position of some of the principal localities and sections using the numbering system followed in the text. HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 7

Table 1. Locality numbers used in this study showing the quadran- Table 1 (continued). gle maps and counties for the locality areas, The widely available No. Locality 15’ Quad 7.5’ Quad County New York State Atlas and Gazetteer (1:150,000; DeLorme Map- 27b Hubbard Quarry Genoa Sheldrake Seneca ping) is a useful modern guide to the topo maps and localities 28 Pleasant Valley New Berlin Sherburne Chenango within the state. An asterisk indicates that the section is illustrated 29a Hicks Point Naples Middlesex Ontario 46c Snyder’s Gully Naples Middlesex Yates- in one of the text-figures. Localities with letter abbreviations of Ontario 15’ quads and numbers are U. S. G. S. localities from de Witt & A-23b Genundewa Point Naples Middlesex Yates Colton (1978). (Nap-4) B Shuman Cemetary Phelps Rushville Yates No. Locality 15’ Quad 7.5’ Quad County (Ph-5) TULLY FORMATION C Sartwell Ravine Penn Yan Penn Yan Yates 1 Beards Creek Caledonia Leicester Livingston (Py-11) 2 Gorham Phelps Rushville Ontario D Lakeside Country Penn Yan Penn Yan Yates 3 Lodi Glen Ovid Lodi Seneca Club (Py-11a) 4 Grove’s Creek Qry Genoa Sheldrake Seneca E Willow Grove Penn Yan Keuka Park Yates 5* Carpenter Falls Skaneateles Spafford Cayuga Crk (Py-14) 6 Borodino Skaneateles Spafford Onondaga F Sunset Point Penn Yan Keuka Park Yates 7* June’s Quarry Tully Tully Onondaga (Py-16) 8* Tinkers Falls Tully Tully Cortland F1 0.9 mi South Penn Yan Keuka Park Yates 9a New Lisbon Hartwick Morris Otsego Crosby 9b Laurens Hartwick Mt. Vision Otsego G Gully N of Ovid Dundee Yates 9c Lebanon Morrisville West Eaton Madison Plum Pt (Ov-3a) H-25a N of Baskin Pt Ovid Dundee Yates GENESEE GROUP (Ov-4) 10* Lake Erie Silver Creek Angola Erie I S of Baskin Pt Ovid Dundee Yates 11* Eighteenmile Crk Eden Eden Erie (Ov-5) 12* Cazenovia Creek Depew Orchard Park Erie J N side of Fir Tree Watkins Glen Reading Ctr Yates 13* Cayuga Creek Depew East Aurora Erie Pt (Wg-2) 14* Elevenmile Creek Attica Corfu Genesee K Green Pt (Wg-7a) Watkins Glen Reading Ctr Schuyler 14a Murder Creek Attica Alexander Genesee L Mill Creek (Ov-10) Ovid Dundee-Lodi Seneca 15* Linden Batavia Batavia South Genesee M N of Tommy Ovid Ovid Seneca 15a* Linden Falls Batavia Batavia South Genesee Creek (Ov-13) 16* White Creek Batavia Stafford Genesee N-27a Romulus Town Ovid Ovid Seneca 17* Taunton Gully Caledonia Leicester Livingston Quarry (Ov-16) 17a Spezzano Caledonia Leicester Livingston O-27b Hubbard Quarry Genoa Sheldrake Seneca 18* Beards Creek Caledonia Leicester Livingston (Gen-2) 18a* Beards Creek Caledonia Leicester Livingston P Ovid-Shedrake Genoa Sheldrake Seneca 19* Fall Brook- Caledonia Geneseo Livingston Road (Gen-1a) Dewey Hill Q Glenwood Crk (I-1) Ithaca Ithaca West Tompkins 19a* Fall Brook Caledonia Geneseo Livingston R Williams Brook Ithaca Ithaca West Tompkins 20* Hemlock Honeoye Honeoye Livingston (I-2) (Big Tree Rd) S S of Williams Ithaca Ithaca West Tompkins 20a Hemlock Honeoye Honeoye Ontario Brook (I-2a) (Huff-Barnard Rd) T Gulf Creek (I-11d) Genoa Ludlowville Tompkins 21 Abbey Gulf Honeoye- Honeoye- Ontario U Esty Glen (I-11-b) Ithaca Ithaca West Tompkins Canandaigua Bristol Center V Twin Glens at Ithaca Ithaca West Tompkins 22 Barnes Gully Canandaigua Canandaigua- Ontario McKinneys (I-11a) Lake W S of McKinneys Ithaca Ithaca West Tompkins 23* Seneca Point Creek Naples Middlesex Ontario (I-11c) 23a Seneca Point Creek Naples- Middlesex- Ontario X Fall Creek, Dryden Ithaca East Tompkins Canandaigua Canandaigua-Lake Ithaca (Dy-10) 23b Genundewa Point Naples Middlesex Yates Y Baker Quarry Dryden Ithaca East Tompkins 24 Bethany Center Batavia Batavia So. Genesee (Dy-10a) (embankment) Z Cascadilla Creek Dryden Ithaca East Tompkins 24b Bethany Center Batavia Batavia So. Genesee (Dy-10b) (roadcut) AA South Hill, Ithaca Ithaca West Tompkins 25 Fir Tree Point Watkins Reading Ctr Yates Ithaca (Dy-10c) (Wg-2-3) BB Enfield Glen Ithaca Ithaca West Tompkins 25a Baskin Pt (Ov-4) Ovid Dundee Yates (I-8a) 26a* Whetstone Brook Honeoye Honeoye Ontario CC Big Salmon Crk, Genoa Genoa Cayuga 27 Lodi Glen Ovid Lodi Seneca Genoa (Gen-13a) 27a Romulus Town Ovid Ovid Seneca DD Big Salmon Crk Genoa Genoa Cayuga Quarry Venice Ctr (Gen-14a) 8 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

Table 1 (continued). Table 1 (continued). No. Locality 15’ Quad 7.5’ Quad County No. Locality 15’ Quad 7.5’ Quad County DD1 Cornwall Gully Penn Yan Penn Yan Yates 47c Lee Cemetery Naples Middlesex Yates (Py-10) 47d School No. 7, Naples Middlesex Yates EE Willow Grove Penn Yan Keuka Park Yates Middlesex (PY-14) 47e Lower East Hill Rd Naples Middlesex Yates FF Sunset Point Penn Yan Keuka Park Yates 47f School No. 2, Naples Middlesex Yates Gully (Py16) Italy Valley GG Creek E of Ovid Dundee Schuyler 48* Wagener Glen Penn Yan Pulteney Steuben Starkey (Ov-6) 49 Bobell Hill Oxford Brisben Chenango HH Granger Point Naples Middlesex Ontario (Nap-3) WEST FALLS GROUP II(XX) 0.8 mi S of Middle- Naples Middlesex Yates 50 Sturgeon Point Silver Creek Angola Erie sex (Nap-8) 51 Big Sister Creek Silver Creek Angola Erie JJ Chidsey Point Penn Yan Pulteney Steuben 52 Eighteenmile Creek Eden Eden Erie Creek (Py-7) 53 Hampton Brook Eden Hamburg Erie KK-F Sunset Point Penn Yan Keuka Park Yates 54* Cazenovia Creek Springville Colden Erie Gully (Py-16) 55* Hunter Creek Depew East Aurora Erie LL-L Mill Creek (Ov-10) Ovid Lodi Seneca 56 Sheldon Creek Arcade Strykersville Wyoming MM Sheldrake Creek Ovid Ovid Seneca 57* Varysburg Attica Attica Wyoming (Gen-1a) 58* Johnson Creek Attica Attica Wyoming NN Coy Glen (I-3) Ithaca Ithaca West Tompkins 59a* Tannery Brook Attica Attica Wyoming OO Lick Brook (I-8) Ithaca Ithaca West Tompkins 60* Relyea Creek Portage Warsaw Wyoming 61* Stony Creek Portage Warsaw Wyoming SONYEA GROUP 63* Kennedy Gulf Batavia Dale Wyoming 30* Eighteenmile Crk Eden Eden Erie 64 Harford Mills Harford Harford Tioga 30a Pike Creek Silver Crk & Angola & Erie 65a* Parrish Gully Naples Middlesex Yates Eden Eden 65* Buck Run Creek Nunda Mount Morris Livingston 31 Smoke Creek Buffalo Buffalo SE Erie 66 Hungerford Quarry Dryden Ithaca East Tompkins 32* Cazenovia Crk Depew Orchard Park Erie 67 Fairfield Forest Dryden Speedsville Tioga 33* Cayuga Creek Attica Cowlesville Erie & 68 Bald Mtn Dryden Speedsville Tompkins Wyoming 69 Owego Owego Owego Tioga 34* Murder Creek Attica Alexander Genesee 70 Elmira Elmira Elmira Chemung 35* Little Tona- Batavia Batavia South Genesee 71* Lake Erie Shore Silver Creek Angola Erie wanda Creek (Farnham Creek) 37 Wyoming Gulf Batavia Wyoming Wyoming 72* Point Breeze Silver Creek Angola Erie 38* Beards Creek Caledonia Leicester Livingston 73* Big Sister Creek Silver Creek Angola Erie 39* Buck Run Creek Nunda Sonyea & Livingston (Angola) Mt Morris 74* Hampton Brook Eden Hamburg Erie 40* Cottonwood Wayland Conesus Livingston 75* Cazenovia Creek Springville Colden Erie Point Gully 77* Glade Creek Arcade Strykersville Wyoming 41 Shurtleff’s Gully Honeoye Livonia Livingston 78* Sheldon Creek Arcade Strykersville Erie & 41a North McMillan Wayland Conesus Livingston Wyoming Creek 79* Varysburg Attica Attica Wyoming 42* Whetstone Brk Honeoye Honeoye Ontario 80 Varysburg Attica Attica Wyoming 43* Briggs Gully Wayland & Springwater & Ontario (Gassman Road) Naples Bristol Sprs 81* Johnson Creek Attica Attica Wyoming 44* Randall Gully Canandaigua Bristol Center Ontario 82* Relyea Creek Portage Warsaw Wyoming 44a Barnes Gully Canandaigua Canandaigua Ontario 83* Stony Creek Portage Warsaw Wyoming Lake 84* Kennedy Gulf Batavia Dale Wyoming 45 Hicks Point Naples Middlesex Ontario 85 Wolf Creek Portage Portageville Wyoming 46 Cook Point Naples Middlesex Ontario 86* Genesee Gorge Portage Portageville Wyoming & 46a Long Point Naples Middlesex Ontario Livingston 46b Whiskey Point Naples Middlesex Ontario 87 Stony Brook Glen Wayland Dansville Steuben 46c Snyder’s Gully Naples Middlesex Yates & 87a I-390–Dansville Wayland Dansville Steuben Ontario 88 Eden Valley Eden Hamburg Erie 46d Griesa Hill Road Naples Middlesex Ontario 89a* Walnut Creek Silver Creek Silver Creek Chautauqua 46e Bristol Springs Rd Naples Bristol Sprs Ontario 89* Walnut Creek Silver Creek Silver Creek Chautauqua 46f Bristol Springs Naples Bristol Sprs Ontario 90* Irish Gulf Eden & Hamburg & Erie Road Gully Springville Colden 47* Conklin Gully Naples Middlesex Yates 91* Glade Creek Arcade Strykersville Wyoming 47a Rumpus Hill Naples Middlesex Yates 92* Beaver Meadow Arcade Strykersville Wyoming 47b Clark Gully Naples Middlesex Yates Creek (Java) HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 9

Table 1 (continued). Table 1 (continued). No. Locality 15’ Quad 7.5’ Quad County No. Locality 15’ Quad 7.5’ Quad County 93 Wiscoy Creek Portage Portageville Allegany 103 Cattaraugus Creek Cattaraugus Gowanda Reserva- 94 Bald Hill, Dryden Speedsville Tompkins (tributary on tion Caroline Cat. Indian Res.) 95 N Chemung Waverly Waverly & Chemung 104 Cattaraugus Creek Cattaraugus Gowanda Cattarau- Wellsburg (S Branch) gus 96 Van Etten Waverly Van Etten Chemung 105 Clear Creek Eden North Collins Erie (North Branch), CANADAWAY GROUP Taylor Hollow 97* Corell’s Point Dunkirk Brocton Chautau- 106 Clear Creek Eden N Collins & Erie qua (North Branch), Langford 98* Little Cana- Dunkirk Brocton Chautau- Marshfield daway Creek qua 107 Anthony Gulf Springville Colden Erie 99* Walnut Creek Cherry Creek Forestville Chautau- 108 Gears Creek Springville Holland Erie qua 109 Java Arcade Strykersville Wyoming 100 Smith Mills Cherry Creek Perrysburg & Chautau- Forestville qua CONNEAUT AND CONEWANGO GROUPS 101 Big Indian Creek Silver Creek Farnham Cattarau- 110 Porter Creek, Clymer Sherman Chautau- gus Summerdale qua 102 Little Indian Creek Silver Creek Farnham Cattarau- 111 Howard Cambridge gus Quarries Springs Edinboro N Erie (PA) until it is last seen on the eastern side of Canandaigua Lake formity (Lower-Upper Tully boundary), and post-Tully dis- at Gage Gully. Farther westward, its place is taken by the conformity (Taghanic Onlap of black Geneseo Shale over Leicester Pyrite, a diachronous remanié deposit marking the gray Moscow Member of the Windom Shale) (Brett & coalescence of horizons at the compound unconformity be- Baird, 1982, 1994, 1996, 1997; Baird & Brett, 1986a, b, tween the Hamilton and Genesee Groups: pre-Tully 2001). The detailed lithostratigraphy of the Tully has been disconformity (Taghanic Unconformity), intra-Tully discon- thoroughly described by Heckel in his excellent paper

Text-fig. 5. Diagram showing the units within the Tully Limestone recognized by Heckel (1973). 10 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

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

(1973) (Text-fig. 5), and summarized by Heckel (1997). Gorham (Ontario County) between Seneca and Canan- Building on Heckel’s work, Baird & Brett (2003) and Baird daigua lakes (Loc. 2), the apparent easternmost occurrence et al. (2003) have added significant detail to the correlations of the Leicester Pyrite, indeterminate tornoceratid nuclei and faunal relations within the Tully, particularly in the occur in pyritic nodules at the sharp contact between the Sherburne-Oneonta area of Otsego and Chenango counties. Bellona Coral Bed of the Tully and the black Geneseo Shale Goniatites are known from a single horizon in the (House, 1965: 83). West of Canandaigua Lake, where the Moravia Bed of the Upper Tully and from black shale inter- Tully is missing, the tornoceratids in the Leicester Pyrite at calations in the Filmore Glen Bed of the uppermost Tully. the base of the black Geneseo Shale that are referable to The horizon in the Moravia Bed is in the upper part of Bed Tornoceras uniangulare uniangulare (Conrad, 1842) were G of West Brook Member of Cooper & Williams (1935: probably all exhumed as pyrite debris from the underlying 791) at Tinkers Falls and June’s Quarry, near Tully (House, Moscow Shale (Baird & Brett, 1986a, b). 1968: 1065) and at corresponding levels farther west (Text- Localities.–Locality numbers with asterisks have sections il- fig. 6). The type locality for Pharciceras amplexum (Hall, lustrated in Text-fig. 6. The one goniatite horizon found 1886: pl. 127 (12), fig. 1) was given as Lodi Landing, Seneca within the Tully Limestone proper is the level yielding Phar- Lake. J. W. Wells located a probable in situ representative of ciceras amplexum. Also listed here are some localities of the this form 0.38 m (1.25 ft) above the Bellona Coral Bed, in Leicester Pyrite (Loc. 1) or localities where goniatites are Lodi Glen, at the same horizon within the Upper Tully as at known close to the Tully (Locs 2 and 4) or, in the east, where Tully and Tinkers Falls. Apart from P. amplexum, tornocer- goniatites apparently occur in equivalents of the Tully (Loc. atids occur at these localities and some have been referred to 9). Detailed correlation was given by Heckel (1973, 1997) Tornoceras cf. arcuatum House (1965: 108). for units of the Tully (Text-fig. 5); see also Baird & Brett Indeterminate crushed molds of probable pharciceratids (2003) and Baird et al. (2003). are known from the black shale interbeds of the Fillmore Locality 1, Beards Creek.–The Leicester Pyrite occurs at Glen Member of the Upper Tully between Skaneateles and the base of the section illustrated in Text-fig. 8 (Loc. 18) at Cayuga lakes. Westward from Cayuga Lake the Upper Tully Beards Creek, Leicester (Livingston County), 0.1 mi (0.16 units are progressively removed by erosional beveling (Text- km) west of the railroad crossing, at an altitude of 638 ft figs. 5, 7), and locally the Leicester Pyrite, with tornocer- (194.5 m). The Leicester Pyrite (Text-fig. 8) here forms dis- atids, is found at the Tully-Geneseo contact and, west of continuous lenses of pyrite nodules and steinkerns. The Canandaigua Lake, at the Moscow-Geneseo contact. At pyrite is also exposed in several neighboring creeks. This is HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 11

the source area for the types of Tornoceras uniangulare uni- Locality 9a, New Lisbon.–Goniatites occur in more clas- angulare. tic equivalents of the Tully Limestone near New Lisbon (Ot- Locality 2, Gorham.–Along Flint Creek, at Gorham sego County). In the northern fork of Stony Brook, 1.5 mi (Ontario County), there are exposures of the Tully Lime- (2.4 km) northeast of New Lisbon, crushed goniatites (poor stone, which here is 1.5 m (5 ft) thick. Exposures were well tornoceratids) occur in equivalents of the West Brook Mem- seen in 1959 east of East Swamp Road, south of Gorham, ber of Cooper & Williams (1935) and presumably in the during excavations for draining the swamp where a pyritic vinicity of the projected level of the West Brook in the New level at the top yielded Tornoceras (House, 1965: 83). Lisbon section illustrated by Heckel (1973: fig. 19). Gordon Locality 3, Lodi Glen.–On the eastern side of Seneca Baird (pers. comm., 2002) reported that the West Brook Lake, at Lodi Point Landing (Seneca County), is the locality Shale is deeply covered in the northern tributary but is still given by Hall (1886: pl. 127 (12), fig. 1) as the source of well exposed in the south branch. This is locality 34 of Baird the type material of Pharciceras amplexum. In Lodi Glen, et al. (2003). above the Landing, J. W. Wells located in situ a probable Locality 9b, Laurens.–At approximately 7 mi (11.3 km) Pharciceras, 0.38 m (1.25 ft) above the Bellona Coral Bed southeast of New Lisbon (Otsego County), Pharciceras am- (Heckel, 1973: 40) in the Tully Limestone. The relations plexum occurs (USNM 96545) in the West Brook Member here are as at Tinkers Falls (Loc. 8). and this is the most easterly record of the genus in the state. Locality 4, Grove’s Creek Quarry.–Southeast of Ovid This locality is 1 mi (1.6 km) northeast of Laurens (Otsego (Seneca County) at Grove’s Creek Quarry, pyritic tornocer- County). This is locality 37 of Baird et al. (2003). atids referred to Tornoceras aff. uniangulare occur below the Locality 9c, Lebanon.–There is a single specimen of Tully Limestone (House, 1965: 83) in the upper part of the Pharciceras amplexum recorded from the West Brook Mem- underlying Moscow Shale. This is within the Gorge Gully ber (USNM 96552) in a ravine 0.5 mi (0.8 km) southwest Submember of the Windom Shale (Brett & Baird, 1994). of Lebanon (Madison County). Locality 5*, Carpenter Falls.–At Carpenter Falls, below the bridge on Appletree Point Road over Bear Swamp Creek, Other Localities.–In this and subsequent entries localities 1 mi (1.6 km) north of New Hope (Cayuga County), the refer to those given in previous publications and follow the level of the Bellona Coral Bed is 0.48 m (1.6 ft) below the previous abbreviations. bridge foundation and the top of the unit above is the ex- Locality Gen-1.–From the black shales of the Fillmore pected level for Pharciceras (Text-fig. 6, Loc. 5). Glen Bed of the uppermost Tully, at Sheldrake Creek, above Locality 6, Borodino.–Tornoceras is recorded from the Cayuga Lake (western side), Gordon Baird and Carlton West Brook Member of the Tully (House, 1965: 83) at Brett collected several molds with well-preserved growth Borodino (Onondaga County; Heckel, 1973: 45). We have lines of probable Pharciceras, one of which preserves ventro- not located the horizon in situ. It is probable that this is from lateral furrows and a flattened venter but it does not show a the same level as other Tully goniatite faunas. suture. De Witt & Colton (1978: locality Gen-1, table 1, pl. Locality 7*, June’s Quarry.–One mi (1.6 km) northeast 3) recorded the base of the Tully at 604 ft (184.1 m) eleva- of Tully (Onondaga County), June’s Quarry (or Ravine) has tion, which is above Blew Road west of NY Rte. 89, approx- provided by far the richest collection from the Pharciceras imately 1.5 mi (2.4 km) southwest of Sheldrake (Seneca level (Text-fig. 6, Loc. 7). Determinable material was first County). found here by W. A. Oliver, Jr. The best collecting is from There are collections at the Paleontological Research In- the old quarry surface on the western side, in the middle stitution (PRI) of similar specimens of probable Pharciceras part of the Moravia Bed of Heckel (1973: 41), 1.27 m (4.2 from the Fillmore Glen Beds at Mack Creek (Loc. Gen-1c), ft) above the base of Bed G of the West Brook Member of near Yarnell Road, 2.6 mi (4 km) northeast of Ovid (Seneca Cooper & Williams (1935). It should be noted that the County; locality 421 of J. W.Wells), at Fernbank (Loc. Gen- Pharciceras level is not in the Apulia Member of Cooper & 5a), along the western shore of Cayuga Lake, 0.5 mi (0.8 Williams (1935), as originally thought (House, 1962: 274). km) north of Taughannock Creek (Tompkins County; local- Locality 8*, Tinkers Falls.–Tinkers or Tinker Falls (Cort- ity 106c of J. W. Wells, Cornell University Museum of Pa- land County), 5 mi (8 km) east of Tully, is the type section leontology [CU] G-89-92); section illustrated by Heckel for the divisions of Cooper & Williams (1935: 791). Here (1973: pl. 2), de Witt & Colton (1978: pl. 3) and Grasso et the Pharciceras bed was located within the Moravia Bed of al. (1986: fig. 8), and Brett & Ver Straeten (1997: 250–251, Heckel, approximately 1.47 m (4.8 ft) above the base of the fig. 35). West Brook Member, in Bed G of Cooper & Williams (Text-fig. 6, Loc. 8), at the same level as in June’s Quarry. 12 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

Text-fig. 7. Facies relations of upper Moscow Shale to lower West River Shale. Alignment of key horizons and the ammonoid and conodont sequences in the upper Moscow and Tully Formations and lower and middle Genesee Group, in central and western New York. Principal remanié 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 after Klapper (1989), Klapper & Johnson (1990), Ziegler & Sandberg (1990), and Klapper & Becker (1999). The change in age of the Leicester Pyrite westward was first demonstrated by Huddle (1974, 1981).

GENESEE GROUP tion and lower Geneseo Shale, and the earlier assignment is This group shows a spectacular thickening eastward thought to be the more probable. Eastward from from 2.7 m (9 ft) at Lake Erie, nearly 40 m (128 ft) in the Canandaigua Lake, the Genesee Group disconformably Genesee Valley, 274 m (900 ft) at Cayuga Lake (de Witt & overlies units of the Tully Formation (Heckel, 1973: pl. 2). Colton, 1978), and perhaps as much as 490 m (1,600 ft) in As presently interpreted, the Genesee Group comprises the Gilboa-Kaaterskill area (Rickard, 1975). In the west, be- beds between the base of the black Geneseo Shale and the tween Lake Erie and Canandaigua Lake, the Genesee Group base of the black Middlesex Shale (Text-fig. 3). The history rests directly on the Moscow Shale of the Hamilton Group of the use of these terms is complex. Vanuxem (1842: 168) (Text-figs 3, 7). At the disconformable junction of the two first used the term Genesee Slate, and his definition at groups, the Leicester Pyrite, containing lenses of nodule- Cayuga Lake indicates that he restricted it to what is now and steinkern-bearing detrital pyrite, is observed. It is a bed called the Geneseo Shale (a term which dates from Chad- named for exposures in the Genesee Valley at Leicester (Liv- wick, 1920). Vanuxem indicated that the Genesee Slate was ingston County). Huddle (1974, 1981) has demonstrated the “Black Shale and Slate, and Upper Black Slate of Re- from conodonts that the Leicester Pyrite becomes progres- ports,” indicating how uncertain was the use of these terms. sively younger in a westward direction. There is strong evi- The present-day practice extends the use of the Genesee ei- dence that the Leicester is a lag, or remanié, unit that ther in the form of a Formation (de Witt & Colton, 1959, includes exhumed material from the Moscow Shale (Baird 1978) or as a Group (Chadwick, 1933; Rickard, 1975), the & Brett, 1986a, b). The rich fauna of Tornoceras uniangulare limits embracing the sequence from the Geneseo Shale uniangulare from the Leicester Pyrite (House, 1965) cannot through the West River Shale, inclusive. This usage essen- unequivocally be assigned to the Moscow Shale, but it is dis- tially dates from Clarke & Luther (1904). The various units similar from tornoceratid faunas known in the Tully Forma- and key marker-beds within have been discussed in detail HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 13

by Brett & Baird (1982, 1986, 1994, 1997), Baird & Brett gasser, 2002). (1986a, b), de Witt & Colton (1959, 1978), Baird et al. (1989), House & Kirchgasser (1993), Kirchgasser (1975, Geneseo Shale, Penn Yan Shale and Associated Units 1985, 1994), Kirchgasser & House (1981), Kirchgasser et In the type area of the Genesee Valley, the Geneseo Shale al. (1986, 1994), Kirchgasser, Brett, & Baird (1997), and (Member) consists of two major tongues of black shale sep- Kirchgasser, Over & Woodrow (1994, 1997). In a series of arated by a sequence of dark gray shales (Text-fig. 8, Locs papers (references above) modeled on their work on the un- 17-19; de Witt & Colton, 1978: pls 2, 6). The lower, or derlying Hamilton Group (Brett & Baird, 1996), Carlton basal tongue, is a continuous sequence of black shale, 1.2- Brett and Gordon Baird described the role of submarine ero- 1.5 m (4-5 ft) thick and overlain by 5.2-6.7 m (17-22 ft) of sion and sedimentary hiatuses associated with eustatic and dark gray shale. tectonic cyclicity in the formation of the black shale, pyrite, The succeeding upper tongue consists of a 1.2-2.1 m (4- and carbonate marker-beds of the Genesee Group. These 7 ft)-thick interval with a pair of black shales, separated by horizons are the keys to correlations within the eastward gray shale, which defines the top of the Geneseo Shale. The thickening and coarsening facies of the group. Many of upper tongue thins westward to a single black shale horizon them are goniatite-bearing beds or proximal to goniatite- at Cayuga Creek (Text-fig. 8, Loc. 13). Thus defined, the bearing beds, which makes them crucial to determining the Geneseo Shale thickens from 1.1 m (3.5 ft) at Cayuga Creek goniatite sequence within the group (see details in the lo- to approximately 9 m (30 ft) in the Genesee Valley (Text-fig. cality descriptions below). 8, Locs 17-19). In approximately the middle of the gray In westernmost sections, west of Cazenovia Creek (Text- shale interval between the lower and upper tongues of the fig. 8, Loc. 12), the Leicester Pyrite is missing and the North Geneseo Shale in the Genesee Valley, there is a prominent Evans Limestone (Rickard, 1964) or “Conodont Bed” of 0.18-0.20 m (7-8 in)-thick argillaceous limestone contain- Hinde (1879), a 75-100 mm (3-4 in) calc-arenitic bone-bed ing crushed Ponticeras. This level, earlier mistaken for the occupies the interval between the upper Moscow Shale of Lodi Limestone (Kirchgasser, 1975), is informally named the Hamilton Group and the overlying upper Genundewa herein the Geneseo Limestone Horizon. It can be traced Limestone of the middle Genesee Group or Formation westward, where the upper part of the gray shale interval ap- along the great Taghanic Unconformity (Text-fig. 7). Traces pears to have been progressively eroded, to Cayuga Creek of the horizon are found beneath the upper Genundewa where its position is in the gray shale beneath the upper Limestone as far east as the Genesee Valley (Kirchgasser, tongue of the Geneseo Shale. 1996a, 1998; Kirchgasser & Koslowski, 1996; Kirchgasser East of the Genesee Valley, the position of the Geneseo & Vargo, 1998). Limestone Horizon is difficult to recognize as the gray shale The North Evans is a winnowed remanié deposit (Baird interval pinches out and the lower and upper tongues of the & Brett, 1986a, b) famous for its abundant fish debris (Hus- Geneseo Shale cannot be differentiated in the eastward sakof & Bryant, 1918; Turner, 1998). Moreover, it is the thickening succession of mostly continuous black shale with famed “conodont bed” of Hinde (1897). Whereas the unit scattered concretionary (septarian) horizons (Text-fig. 3; de at its type area at Eighteenmile Creek (Loc. 11) (Text-fig. Witt & Colton, 1978: pls 2-3, 5). At Canandaigua Lake, 8) appears to occupy the time-rock interval between the the Geneseo Shale is 13 m (44 ft) thick at Menteth Gully (de upper Givetian varcus Zone and lower Frasnian MN (Mon- Witt & Colton, 1978). Farther east, at Seneca and Cayuga tagne Noire) Zone 2 (Text-fig. 7), recent taphonomic work lakes, Baird & Brett (1986b) and Baird et al. (1989) recog- (including that of Kirchgasser, 2001, 2004) suggests that nized the Fir Tree Limestone (and equivalent Fir Tree Pyrite) most of the microvertebrate and conodont elements in the as a submember in the upper Geneseo Shale. The Fir Tree North Evans, throughout its geographic extent, are of lower Limestone occurs at the base of the Hubbard Quarry Shale, Frasnian (MN Zone 2) age, including the youngest zonal a submember of dark gray shale, comprising the uppermost indicator in the unit, Ancyrodella recta Kralick, 1994; MN division of the Geneseo Shale (Text-fig. 7). At the spectac- Zone 3 begins in the overlying upper Genundewa Lime- ular Taughannock Falls on the western side of Cayuga Lake, stone with the entry of an early form of A. rugosa Branson the Geneseo Shale is nearly 60 m (183 ft) thick and its top & Mehl, 1934. Recently, pyritic protoconchs and early lies just below the lip of the falls (Baird & Brett, 1986b: 60- whorls of probable Frasnian goniatites (?Koenenites) have 61, fig. 12). been recovered from the North Evans, which supports the Goniatites are poorly known in the Geneseo Shale and view that the unit mostly records a sub-upper Genundewa most specimens are crushed flat with the shells having been disconformity with parts of the upper Penn Yan Shale and removed in the dysoxic to anoxic basinal environments of nodular lower Genundewa reworked or removed (Kirch- deposition. These include rare Pharciceras? sp. at localities 14 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

Text-fig. 8. Sections of the lower Genesee Group comprising the Geneseo Shale, Penn Yan Shale, and Genundewa Limestone between Lake Erie and Hemlock Lake, with the locality numbers used in the text. For Loc. 19/6 (LH) read barite (for baryte). HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 15 16 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

in Chenango and Otsego counties. Tornoceras sp. occurs side of Seneca Lake, a Lodi horizon (Lodi Bed C), 5.5 m abundantly at 1.4 m (4.5 ft) (Text-fig. 8, Bed 18/1) and 5.1 (18 ft) above the main Lodi Bed B, yielded the large speci- m (16.75 ft) (Bed 18/2) above the Leicester Pyrite at Beards men of P. cf. perlatum figured herein (Text-fig. 26D-E, Loc. Creek (Loc. 18) in the Genesee Valley, but only one speci- M, Ov-13). The Lodi and post-Lodi horizons are the prob- men is known with a sharply projecting ventrolateral furrow able source of the records of P.perlatum, many in beds with that could suggest a pharciceratid (from Bed 18/1). The oc- aulaporid corals (Cladochonus beds or zones), in the Sher- currence of Ponticeras in the Geneseo Limestone Horizon of burne Siltstone localities of Kindle (1896) and Boekenkamp the Genesee Valley has been noted above. Epitornoceras en- (1963) at the south end of Cayuga Lake (Locs I-1, I-11a–d). ters in the succession, as E. cf. mithracoides (Frech, 1887), Westward (basinward) from Seneca Lake, the Lodi beds approximately 12.2 m (40 ft) below the top of the Geneseo converge to a condensed sequence of reworked pyrite-phos- Shale at Lodi Glen, Seneca Lake (Loc. 27) (House, 1965: phate lags beneath black shales. In the Honeoye Lake Valley, 119), 24-27 m (80-90 ft) above the top of the Tully Forma- on the western side of the regional basin axis, Lodi Bed B tion. A specimen of E. cf. peracutum (Hall, 1879) comes reappears as a single auloporid-bearing carbonate horizon from near the top of the Geneseo Member at Hubbard capped by a black shale designated Penn Yan Black Shale A, Quarry (Loc. 27b) (House 1965: 119) on the western side a horizon which marks the conodont Middle/Upper Devon- of the Cayuga Lake valley. The holotypes of E. peracutum ian boundary in the region (Baird & Brett, 1986b; Baird et and E. aff. peracutum (see House, 1978: 60) are from the al., 1989; Kirchgasser et al., 1989; Kirchgasser, 1994). A sec- Ithaca Formation. Pyritic fragments and protoconchs of go- ond black shale (Penn Yan Black Shale B), approximately 2 niatites (Tornoceras?, Ponticeras?) have been noted by us in m (6 ft) above the Lodi at Abbey Gulf (Loc. 21) in the Ho- the residual pyrite of the Fir Tree Limestone in the Salmon neoye Valley, has yielded Ponticeras. Creek Valley east of Cayuga Lake (Locs CC, Gen-13a; DD, Farther to the west, in the Genesee Valley, the Lodi 14a). Ponticeras perlatum (Hall, 1874) enters the succession Limestone is represented by a single bed of nodules, with in the dark gray to black shales at the top of the Geneseo Ponticeras perlatum, in the lower Penn Yan Shale approxi- Shale (Hubbard Quarry Shale), the best locality being Hub- mately 1 m (3 ft) above the top of the Geneseo Shale (Text- bard Quarry (Loc. 27b), which has yielded many specimens, fig. 8, Loc. 18). Immediately above the Lodi is a pair of thin although most are badly crushed; rarely are they in vertical black shales that represent Penn Yan Black Shales A and B. orientation (Pl. 4, Fig. 5). These post-Lodi black shales converge westward to a single black shale unit, thus constraining the top of the Lodi as far Penn Yan Shale and its Eastern Equivalents west as Cayuga Creek where the Lodi horizon is represented The Lodi Limestone (Lincoln, 1895), exposed in Lodi Glen by a thin, grey mudstone (Text-fig. 8, Loc. 13). on Seneca Lake (Loc. 27), is a 0.23 m (9 in)-thick concre- A thin black shale layer (Schumacher Bed [SB]), overly- tionary bed 1.2 m (4 ft) above the top of the Geneseo Shale. ing a concretion band 4-5 m (12-17 ft) above the Lodi hori- The Lodi Limestone lies in the lower part of the local Penn zon in the Genesee Valley, is a key horizon for positioning Yan Shale (or Sherburne Siltstone). At this locality, at the goniatite occurrences higher in the Penn Yan Shale and Romulus Town Quarry, Ovid (Loc. 27a), and at Hubbard equivalent Sherburne Siltstone and the Ithaca Shale and Quarry (Loc. 27b) above Cayuga Lake, the Lodi bed has Sandstone. The SB black shale (Kirchgasser et al., 1986: yielded fine specimens of Ponticeras perlatum, the type of 249) and underlying concretion bed have been traced west- which came from Homer (Cortland County), farther to the ward to Cayuga Creek (Text-fig. 8, Loc. 13). It is the posi- east (House, 1965; Kirchgasser, 1975). tion of the SB black shale at Honeoye and Canandaigua The main bed of the Lodi Limestone at Lodi Glen is one lakes that constrains the position, in western sections, of the of several recurring horizons of impure carbonates with aulo- Renwick Shale, a major black shale marking the top of the porid corals and other benthic fossils in the eastward thick- Sherburne Siltstone and base of the Ithaca Shale and Sand- ening wedge of Penn Yan or Sherburne facies in the Seneca, stone in the Seneca and Cayuga valleys. At Abbey Gulf (Loc. Cayuga, and Skaneateles valleys (Baird & Brett, 1986a, b; 21) in the Honeoye Lake Valley, the western tongue of the Baird et al., 1989). On the western side of Seneca Lake these Sherburne Siltstone, the Renwick Shale, and the informal recurring Lodi beds, designated Lodi Beds A-D, and similar unpublished “Abbey Gulf Beds” of G. Baird and C. Brett horizons higher up, can be seen in the bluff exposures (Locs (Baird et al., 2006: 355; Zambito et al., 2007: 94, fig. 4), 25, 25a, Ov-5, Wg-2, Wg-7a); Lodi Bed B is the main Lodi with indeterminate goniatites, form a sequence in the nearly bed and Lodi Bed A is at the Geneseo/Penn Yan-Sherburne 9 m (30 ft) interval between the Lodi Limestone (Lodi Bed contact. Some of the horizons have yielded Ponticeras but B) and the SB black shale. Thus, the many records of Pon- they have not been systematically searched. On the eastern ticeras perlatum reported from the top of the Sherburne and HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 17

base of the overlying Renwick Shale at localities at the south 1985) could conceivably be the Crosby Horizon. In the Lick end of Cayuga Lake are stratigraphically below the SB black Brook section (Loc. I-8) of de Witt & Colton (1978: pl. 3), shale. These are the highest confirmed records of P. perla- the Bluestone-Crosby bed would be the siltstone unit im- tum. mediately below the covered interval in approximately the Higher in the Cayuga Lake Genesee Formation succes- middle of the Ithaca. sion, in the lower third of the Ithaca Shale and Sandstone, Acanthoclymenia enters the succession in the upper Penn Chutoceras nundaium (Hall, 1874) occurs in the Quarry Yan in Bed 15/9 on top of the Linden Horizon at Linden Sandstone interval at Fall Creek, Ithaca (Loc. Dy-10), 72 m (Genesee County; Loc. 15) and there is also a record in the (235 ft) above the base of the Renwick Shale. It also occurs uppermost Penn Yan Shale at Sunset Point, Keuka Lake in the same interval at South Hill, Ithaca (Loc. Dy-10c) (Loc. F, Py-16), 0.30 m below the Crosby Sandstone. (Kirchgasser, 1985), and possibly also in the Williams Brook Coquinite at Williams Brook (Loc. I-2) on the southwestern Localities.–Spectacular eastward thickening is shown by the side of Cayuga Lake. Unfortunately, none of these horizons pre-Genundewa Genesee Group. On the accompanying di- has been found to the west of Cayuga Lake, although C. agrams (Text-fig. 8), therefore, different scales are used for nundaium is represented in the equivalents of the upper the columns. The overlying Genundewa Limestone, which Penn Yan Shale at Mill Creek, Seneca Lake (Loc. L, Ov-10). is thin in relation to the lower levels, is also shown at an in- Koenenites enters the succession in the thin styliolinid creased scale for clarity; the Genundewa is informally sub- limestones that characterize the upper Penn Yan Shale. The divided into a nodular-bedded lower Genundewa and a most productive horizon is the Linden Horizon (Kirchgasser well-bedded upper Genundewa. The nodular-bedded lower & House, 1981) in which Koenenites styliophilus styliophilus Genundewa is missing in western sections between Linden (Clarke, 1898) is locally common and can be well preserved. (Loc. 15) and Lake Erie (Loc. 10). Locality numbers with as- This horizon has been traced from Cayuga Creek (Loc. 13) terisks have sections illustrated in Text-figs 8-9. in the west to the section above the Quarry Sandstone inter- Locality 10*, Lake Erie.–On the shore of Lake Erie, there val at Fall Creek (Loc. Dy-10), Ithaca, at Cayuga Lake is a complete section of the Genesee Group, from the upper- (Kirchgasser, 1985). most Windom Member of the Moscow Shale (Hamilton Koenenites styliophilus styliophilus is also locally common Group) to the base of the Middlesex Black Shale (Sonyea and well preserved in the Crosby Sandstone of Torrey et al. Group). This section is seen in the cliffs 170 yd (155 m) (1932) in sections in the vicinity of Keuka Lake (Locs C, south of the mouth of Pike Creek, 1.4 mi (2.25 km) north- Py-11, D, Py-11a, F, Py-16). At Keuka Lake, the Crosby east of Jerusalem Corners (Erie County; Text-fig. 8, Loc. Sandstone marks the contact between the Penn Yan Shale 10). No goniatites have been collected there. A 31 mm (1.25 and the overlying Ithaca Shale and Sandstone. To the west in) unit at the base consists of laterally connected calcarenitic at Seneca Point Creek, Canandaigua Lake (Loc. 23), Koe- lenses, 6 mm (0.25 in) thick [North Evans Limestone with nenites occurs in the “hiatus concretion horizon” of Baird mm-thick lenses of pyrite (?Leicester Pyrite) at its base; (1976), a bed with the distinctive conodont fauna of the Kirchgasser, 2004], a black shale, 19 mm (0.75 in) thick, Crosby, which is found 7 m (23 ft) below the base of the and a Styliolina-rich bed, 6 mm (0.25 in) thick at the top. Genundewa Limestone. To the east at Mill Creek, Seneca Above this unit is 0.23 m (9 in) of interbedded very dark Lake (Loc. L, Ov-10), the Crosby lies 6.1 m (20 ft) above gray and black shale that represents undifferentiated Penn the Linden Horizon and above other higher horizons also Yan Shale. The upper Genundewa Limestone lies above. A yielding Koenenites. Although the Crosby Sandstone appears nearby section at the mouth of Pike Creek was illustrated to occupy the stratigraphic position of the Genundewa by de Witt & Colton (1978: pl. 2). Limestone east of Canandaigua Lake in some respects (see Locality 11*, Eighteenmile Creek.–Along Eighteenmile de Witt & Colton, 1978: pls 3-5), we believe it is an older Creek, 137 yd (125 m) downstream from the westernmost horizon in the interval between the Linden Horizon and the of two railroad bridges, on the northern side of the creek, Genundewa. The upper Penn Yan Shale appears to be trun- 1.2 mi (1.9 km) northwest of North Evans (Erie County), cated in the region of Keuka Lake and it could be that there there is an even more condensed sequence (Text-fig. 8, Loc. the Crosby Sandstone cuts down to the level of the Linden 11). Here, beneath the upper Genundewa Limestone, is a se- Horizon and incorporates Linden goniatites as a basal de- quence of 6 mm (0.25 in) of black shale beneath 86 mm posit. No goniatites or fossils other than plant fragments are (3.4 in) of North Evans Limestone, followed by 25 mm (1 known from the Crosby farther east. At Fall Creek, Cayuga in) of very dark gray to black shale representing the Penn Lake (Loc. Dy-10), the prominent siltstone immediately Yan Shale. Pyritic specimens of ?Koenenites have recently above the Linden Horizon (Bluestone Bed of Kirchgasser, been noted in microfossil residues from the 29-mm-thick 18 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

Bed 11/1e at the top of the North Evans Limestone (Kirch- lower of the pair of black shales above the Lodi to the east gasser, 2002). No other goniatites are known from this lo- (Penn Yan Black Shale A). It is not the Renwick Shale and cality, which is a classic section of Grabau [1898–1899, top of the Geneseo Shale as previously suggested (Kirch- Section I (H); illustrated by de Witt & Colton (1978: pl. 2) gasser & Brett, in Oliver & Klapper, 1981b: 19, fig. 1; reil- and Kirchgasser (1981: 11, fig. 1, reillustrated by Brett et lustrated by Kirchgasser, Brett, & Baird, 1997: 197, fig. 13). al., 1997: 178, fig. 7)]. Poorly preserved goniatites occur in Bed 13/3, the Locality 12*, Cazenovia Creek.–In Cazenovia Creek, a prominent bed of concretions in the Penn Yan, which is be- similar section is seen, but in different facies. One mile (1.6 lieved to be the concretion horizon below the Schumacher km) south of Spring Brook (Erie County), and below the or SB black shale farther east. Bed 13/4 is the Linden Hori- intersection of North Davis and Conley roads, a westward zon, a styliolinid-rich concretionary bed which farther east flowing tributary joins Cazenovia Creek, and the section is the source of well-preserved Koenenites, but no specimens given here (Text-fig. 8, Loc. 12) is 20 yd (18 m) downstream have been noted here. Bed 13/5 is the bed of grotesque con- of the confluence. A thin lens (6 mm or 0.25 in) of Leicester cretions at the top of the Penn Yan Shale. Immediately above Pyrite is followed there by 0.66 m (2.2 ft) of black and dark the concretion layer, mm- to cm-thick seams of North Evans gray shale (undifferentiated Geneseo and Penn Yan shales) fish-bone and conodont debris encrust the lower surface of with a 25 mm (1 in) bed of fine-grained limestone, 0.20 m the Genundewa Limestone. The reworked fish and con- (8 in) above the base. There are small carbonate concentra- odont debris of the North Evans, associated with the sub- tions near the top of this interval, 76 mm (3 in) below the upper Genundewa unconformity, occurs as far east as upper Genundewa Limestone. Bed 12/3, a 25 mm (1 in)- Taunton Gully (Loc. 17) in the Genesee Valley (Kirchgasser, thick seam of laminated calcarenite on the underside of the 1996a, 1998; Kirchgasser & Koslowski, 1996; Kirchgasser upper Genundewa Limestone, contains the rich conodont & Vargo, 1998). The section was illustrated by de Witt & and fish-bone fauna of the North Evans Limestone (see ref- Colton (1978: pl. 2). A detailed log was given by Brett & erences under Loc. 13). A similar section on the western side Baird (1990: A-19, fig. 6; reillustrated by Kirchgasser, Brett, of the creek, below where the power lines cross Northrup & Baird, 1997: 195, fig. 12). Road, was illustrated by Brett & Baird (1982: 34, fig. 5B) Locality 14*, Elevenmile Creek.–In Elevenmile Creek, and Baird & Brett (1986a: 166, fig. 4A). The section was the section illustrated commences approximately 230 yd also illustrated by de Witt & Colton (1978: pl. 2). (210 m) downstream of a railroad bridge, 1.3 mi (2.1 km) Locality 13*, Cayuga Creek.–The Cayuga Creek section, southwest of Darien Center (Genesee County). Bed 14/2, in 2 mi (3.2 km) northwest of Cowlesville (Wyoming County) the Penn Yan Shale, 1.19 m (3.9 ft) below the upper Ge- shows the entry of some of the characteristics of more east- nundewa, yields goniatites and correlates eastward with the erly successions. The top of the illustrated section (Text-fig. Linden Horizon (Bed 15/8) at Linden (Loc. 15). 8, Loc. 13) is 120 yd (110 m) upstream of the NY Rte. 354 Locality 15*, Linden.–The section near Linden (Genesee bridge, where the Genundewa Limestone forms a falls below County) is in a small, eastward-flowing tributary to Little the top of an old concrete dam. The interpretation of beds Tonawanda Creek, 0.3 mi (0.48 km) north of the bridge at differs in detail from accounts given earlier (see references Linden. The Moscow/Geneseo contact is at the confluence below). As at Cazenovia Creek, the Leicester Pyrite forms a of the tributary and creek. A group of black shales and con- basal marker between the gray Moscow Shale and the black cretion horizons is exposed in a falls section capped by the Geneseo Shale. The 0.28 m (11 in)-thick black shale band Genundewa Limestone (lower and upper parts). The Lodi immediately above the Leicester marks the base of the Gene- Limestone horizon is Bed 15/4, a bed of nodules beneath a seo to the east (lower black shale tongue of de Witt & 0.61 m (2 ft)-thick black shale which is equivalent to the Colton, 1959: 2815). Beds 13/1 and 13/2 are thin black pair of marker black shales traceable above the Lodi eastward limestones in the succeeding 0.65 m (2.2 ft) of dark gray to to the Genesee Valley. Bed 15/5, the most prominent bed of black shale with horizons rich in the diminutive brachiopod concretions in the falls section, underlies a 0.30 m (1 ft)- Devonochonetes and current-oriented Styliolina. Bed 13/2 oc- thick black shale believed to be the SB marker black shale. cupies the position of the Geneseo Limestone farther east. Nearer the top of the Penn Yan Shale, the Linden Horizon The 0.13 m (5 in)-thick black shale band above the interval (Bed 15/8) is expressed as a distinctive bed of nodules, some- of dark gray to black shale is the upper black shale tongue times spherical and pyritic, with Koenenites beneath a 76 (and top) of the Geneseo Shale. Above is a 10 mm (4 in)- mm (3 in)-thick seam of styliolinid limestone (Bed 15/9) thick interval of gray mudstone which is equivalent to the which underlies a 0.18 m (7 in)-thick black shale, the base Lodi Limestone horizon of the lower Penn Yan Shale to the of which is approximately 2.1 m (7 ft) below that of the Ge- east. The overlying 0.23 m (9 in)-thick black shale is the nundewa Limestone. The Linden Horizon, which locally HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 19

yields well preserved K. styliophilus styliophilus, can be traced accessible near the two falls below the bridge where Starr eastward to the Genesee Valley and possibly as far east as the Road crosses the creek near the old Taunton Cemetery. The Keuka, Seneca, and Cayuga Lake valleys (Kirchgasser, Lodi Limestone horizon is a distinctive horizon of small 1985). Shale partings at the base of Bed 15/9 yield crushed nodules (Bed 17/2), 0.84 m (2.75 ft) above a 0.25 m (10 specimens of Acanthoclymenia sp. The section was illustrated in)-thick black shale exposed near the level of the large pool by de Witt & Colton (1978: pl. 2). at the base of the lower falls; the top of the black shale marks Locality 15a, Linden Falls.–A similar sequence is seen in the Geneseo/Penn Yan contact. The Lodi bed locally yields the face of the falls over the Genundewa, below the bridge Ponticeras, and the overlying pair of black shales (the lower at Linden. There Bed 15a/5, the same horizon as Bed 15/5, one is Penn Yan Black Shale A, the upper Black Shale B) lies 0.30 m (1 ft) above the level of the pool at the base of provide useful marker horizons on both sides of the Genesee the falls. Bed 15a/8, the Koenenites-bearing Linden Horizon, Valley. occurs 1.96 m (6.4 ft) above Bed 15a/5. The 20-mm-thick SB marker black shale (Bed 17/2a) Locality 16*, White Creek.–A discontinuous section lies 0.38 m (1.25 ft) above the first prominent bed of con- through the Lower Genesee crops out along the western cretions (with indeterminate goniatites) in the face of the branch of White Creek near farm buildings downstream main falls and 3.8 m (12.6 ft) above the Lodi horizon. Koe- (north) of the NY Rte. 20 overpass, 2.1 mi (3.4 km) east of nenites occurs in Bed 17/3, a horizon of concretions 0.58 m Bethany Center (Genesee County). In this section, the (1.9 ft) above the SB black shale and 1.2 m (4 ft) below a Geneseo Limestone horizon is the resistant 0.15 m (6 in)- still higher concretionary horizon (Bed 17/4) with Koenen- thick limestone which caps a small falls downstream of the ites and Epitornoceras? at the lip of the falls. The main Koe- farm, 3.5 m (11.5 ft) above the Moscow-Genesee contact. nenites horizons (K. styliophilus styliophilus) occur above the This is the westernmost locality where the Geneseo Lime- lip of the falls in the concretions of Bed 17/5 and immedi- stone horizon has been identified in outcrop, but the hori- ately above in the pyritic nodules of the Linden Horizon zon correlates westward into the cluster of three (Bed 17/6). The Linden underlies a 0.13 m (5 in)-thick concretionary bands beneath the base of the upper black black shale and is 5.03 m (16.5 ft) below the base of the Ge- shale tongue of the Geneseo as far west as Cayuga Creek nundewa Limestone, which crops out below the bridge. The (Loc. 13). Above the Geneseo Limestone Horizon, expo- section was illustrated by de Witt & Colton (1978: pls 2, 6). sures are poor, but in an open field below the farm buildings, Locality 17a, Spezzano Gully.–A section similar to that exposures resume with a horizon of large concretions (Bed in the upper part of Taunton Gully is exposed in Spezzano 16/3a) below the pair of marker black shales in the lower Gully west of the railroad crossing, 0.9 mi (1.4 km) farther Penn Yan Shale. Bed 16/3a is in the position of the Lodi north, and 2.6 mi (4.2 km) north of Leicester (Livingston Limestone. Above this level, Ponticeras? occurs in the concre- County). The productive horizons are Bed 17a/3, a horizon tions of Bed 16/4 which lies 1.17m (3.8 ft) above the higher of large concretions 0.91 m (3 ft) above the lip of the main of the two black shales. A 51 mm (2 in)-thick black shale falls, and Bed 17a/4, which is taken as the main Koenenites (?SB black shale) lies 0.61 m (2 ft) above. In the small em- horizon (Linden Horizon), a layer of nodules on top of a bankment on the northern side of the creek, Bed 16/5 is the 0.13 m (5 in)-thick limestone and beneath a 76 mm (3 in)- nodular Linden Horizon with Koenenites (thought to be the thick black shale, 0.96 m (3.2 ft) above Bed 17/a3 and 4.1 same as Beds 15/8 and 15a/8 at Linden) and lies 0.13 m (5 m (13.5 ft) below the Genundewa Limestone. in) below the 51-mm-thick black shale. Bed 16/6 is a 76 Locality 18*, Beards Creek.–Along Beards Creek, 0.3 mi mm (3 in)-thick styliolinid band on top of Bed 16/5. Thin (0.48 km) northwest of Leicester (Livingston County), is de goniatite-bearing styliolinid limestones (Bed 16/6a, 16/7) Witt & Colton’s (1959) reference section for the Genesee are exposed near the culvert under NY Rte. 20. The higher Group. The section commences above the Leicester Pyrite, beds of the Penn Yan Shale are poorly exposed in the creek 0.1 mi (0.16 km) from the railroad crossing of the creek. bed on the south side of NY Rte. 20, below outcrops of the Here a 1.2 m (4 ft)-thick, lower black shale tongue of the Genundewa Limestone. Geneseo Shale is followed by 6.8 m (22.4 ft) of dark gray Locality 17*, Taunton Gully.–A well-exposed succession shale with thin limestones that in turn are overlain by a 0.38 of the lower Genesee is found in Taunton Gully, approxi- m (1.25 ft)-thick upper black shale band. The two black mately 670 yd (612 m) upstream from the railroad crossing shale units respectively represent de Witt & Colton’s (1959) of Pearl Creek, 1.7 mi (2.7 km) north of Leicester (Liv- lower and upper tongues of the Geneseo. The intervening ingston County). Here the Geneseo Limestone (Bed 17/1) dark gray shales were regarded by them as a tongue of Penn is the prominent 0.2 m (8 in)-thick unit 5 m (16.4 ft) above Yan Shale, but they are now included in the Geneseo Shale. the Leicester Pyrite. The higher parts of the section are most Tornoceras is relatively common in Bed 18/1, within the first 20 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

0.3 m (1 ft) above the top of the lower tongue of black shale. The higher part of the Penn Yan can be measured in the Bed 18/2 lies immediately above the highest of five thin side creek, but a more accessible continuation of the succes- black shales in the succeeding 1.8 m (5.8 ft) of the section. sion is seen in the NY Rte. 39-20A roadcut along Dewey The Geneseo Limestone (Bed 18/2a), the prominent Hill, 0.2 mi (0.32 km) to the north. The nodules of Bed limestone in approximately the middle of the gray shale in- 19/4 (Lodi Limestone) and the capping 77 mm (3 in)-thick terval, lies 3.9 m (12.8 ft) above the top of the lower black black shale (Black Shale A) are exposed in the ditch on the shale tongue. Bed 18/3, also with Ponticeras, is a thin lime- southern side of the road where the outcrop begins. The stone 0.35 m (1.2 ft) above. The nodular Lodi Limestone prominent 0.18 m (7 in)-thick black shale (Bed 19/4a), (Bed 18/4) with P. perlatum lies beneath a 76 mm (3 in)- overlying a line of concretions, approximately 5.2 m (17.2 thick black shale (Black Shale A of the Penn Yan Shale), 1.42 ft) above the Lodi, is the SB black shale horizon. Although m (4.6 ft) above the top of the upper black shale tongue of not yet recognized in the Genesee Valley sections, the black the Geneseo Shale and 0.66 m (2.2 ft) above the top of the Renwick Shale and the succeeding Abbey Gulf Beds of more 0.3 m (1 ft)-thick black shale, the top of which defines the eastern sections are believed to project into the interval be- contact between Geneseo and Penn Yan Shales. Ponticeras tween the Lodi and SB horizons. also occurs in an interval of gray shale (Bed 18/5) exposed In the gray shales 1.2 m (4 ft) above the SB horizon is a in the southern side of the creek, downstream of the high prominent layer of large concretions (Bed 19/4b), and 1.8 falls over the Genundewa Limestone below the Dunkley m (5.75 ft) above is Bed 19/5, the first of a series of thin Road bridge. Bed 18/5 is the interval 1.7-4 m (5.6-13.2 ft) limestones with goniatites. Preservation is poor but expo- above the Lodi, and the large septarian nodules (Bed 18/6) sures of Bed 19/5 on the northern side of the road have just above might be the concretion horizon below the SB yielded Ponticeras? and a large tornoceratid (?Epitornoceras). black shale. Horizons higher than Bed 18/6 are not readily Bed 19/5 lies 8.6 m (28.25 ft) above the Lodi horizon. On accessible for collecting. The section was illustrated by de the southern side of the road, Bed 19/6, which lies 0.97 m Witt & Colton (1978: pl. 5). (3.2 ft) above the base of Bed 19/5, is the Linden Horizon, Locality 19*, Fall Brook-Dewey Hill.–On the eastern the first of a series of thin styliolinid beds. Bed 19/6 contains side of the Genesee Valley, the well-known locality noted by small baritic replacements of Koenenites styliophilus stylio- James Hall (1843: 462) at Fall Brook, 1.5 mi (2.4 km) south philus and Tornoceras. Goniatites also occur in the Bed 19/7 of Geneseo (Livingston County), provides a spectacular ex- approximately 0.42 m (1.4 ft) above the base of Bed 19/6 posure from the upper Moscow Shale through the lower and in the highest styliolinid band (Bed 19/8), which is 3.07 Genesee Group. In the cliff above the northern bank of the m (10 ft) above the base of Bed 19/6 and approximately 3 creek downstream of the falls over the Genundewa Lime- m (10 ft) below the base of the Genundewa Limestone, stone, the lower and upper tongues of the Geneseo Shale, which crops out in the bushes at the top of the section. each approximately 1.5 m (5 ft) thick, are separated by 5.2 This section has been illustrated by de Witt & Colton m (17 ft) of dark gray shale. Lenses of the Leicester Pyrite are (1978: pl. 2), Oliver & Klapper (1981b: 37), Kirchgasser et well exposed at the Moscow-Geneseo contact. The promi- al. (1994: 51), and Brett & Ver Straeten (1997: 222). nent 0.17 m (7 in)-thick limestone (Bed 19/3) with crushed Localities 20* and 20a, Hemlock.–The nearest useful ex- Ponticeras, in approximately the middle of the gray shale in- posures of the lower Genesee east of Fall Brook are those terval, is the Geneseo Limestone. This widespread horizon north of Hemlock (Livingston County), almost 13 mi (21 in the Genesee Valley, which there lies 4 m (13 ft) above the km) distant, a fact that makes detailed correlation difficult. base of the lower black shale tongue, was previously referred In the westward-flowing tributary to Hemlock Outlet, to the Lodi Limestone (Kirchgasser, 1975; Kirchgasser & which crosses Big Tree Road 0.9 mi (1.4 km) northeast of House, 1981; Oliver & Klapper, 1981b), but that horizon Hemlock (Loc. 20), the upper part of the Penn Yan Shale is is now known to be higher in the section. well exposed below a falls above the Genundewa Limestone. The section is accessible in a small tributary that enters A a similar setting is found in another tributary (Loc. 20a) Fall Brook at the eastern edge of the cliff. In the side creek, that crosses the Huff-Barnard road intersection 0.9 mi (1.4 the 1.4 m (4.5 ft)-thick upper black shale tongue of the km) farther north. Beds 20/1, 20/2, and 20/3 are thin lime- Geneseo Shale lies 0.20 m (8 in) above a thin limestone (Bed stones with goniatites that crop out within the first 2.7 m (9 19/3a), which caps a falls. The Lodi Limestone is the nodu- ft) above where exposures begin. At Loc. 20a, thin black lar bed (Bed 19/4) in the lower Penn Yan Shale that lies be- shales were noted above equivalents of Beds 20/2 and 20/3. neath a thin black shale (Black Shale A), 2.9 m (9.5 ft) above In the upper part of the main section, Beds 20/4 and 20/5 the base of the upper black shale tongue of the Geneseo form a pair of thin limestones (with poor goniatites) 50-75 Shale. mm (2-3 in) thick and 0.3 m (1 ft) apart. These occur 1.75 HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 21

m (5.75 ft) below a layer of large concretions at the foot of in)-thick black shale taken to be the SB black shale of west- the falls below the Genundewa Limestone. Bed 20/4 (and ern sections, which thus constrains the interval of the Sher- 20a/4) is taken as the Linden Horizon. burne, Renwick, and “Abbey Beds” to the interval between Locality 21, Abbey Gulf.–A well-exposed and almost the Lodi and the SB black shale horizon in the Genesee Val- complete section through the lower Genesee Formation is ley and farther westward (Text-fig. 8). The SB black shale is found in Abbey Gulf, east of Allens Hills Road, 1.6 mi (2.6 the first black shale below sample-horizon 6697 SD in km) northeast of Honeoye (Ontario County; Baird et al., Menteth Gully (de Witt & Colton, 1978: pls 2-3). 1989: figs. 7A, 9B). Although the goniatite record there is Approximately 3 m (10 ft) above the SB black shale, a poor, several marker beds in the section are important for 0.10 m (4 in)-thick limestone, beneath a 0.61 m (2 ft)-thick correlations eastward to Canandaigua Lake and beyond. black shale, caps a low falls. This limestone correlates west- References are made to the sections by de Witt & Colton ward with Bed 20/3 at Hemlock. Cropping out 2.4 m (8 ft) (1978: pl. 2): Cd-Abbey Gulf, Cd-6-Vincent, and Cd-7- above the top of the falls is a pair of thin limestones, Bed Menteth Gully (Canandaigua Lake). At Abbey Gulf and 21/2, 0.10 m (4 in) thick, and Bed 21/3, 76 mm (3 in) nearby sections in the Honeoye, Bristol, and Canandaigua thick, 0.30 m (1 ft) higher. Although no goniatites have Lake valleys, the Middle-Upper Devonian (Givetian-Frasn- been found, Beds 21/2 and 21/3 are taken to be the Linden ian) boundary is located above the Lodi Limestone in the Horizon, which correlates westward to Beds 20/4 and 20/5 lowermost Penn Yan Shale (Kirchgasser et al., 1989; Kirch- at Hemlock and Bed 19/6 at Fall Brook-Dewey Hill (Text- gasser, 1994: fig. 3). fig. 8). At Abbey Gulf, a gas seep reaches the surface in the In the lower part of the Abbey Gulf section, at a level beds of the upper Penn Yan below the falls capped by the taken to be the top of the upper black shale tongue and top Genundewa Limestone. Bed 21/5 is a 76 mm (3 in)-thick of the Geneseo Shale, is the Lodi Limestone (Bed 21/1), a bed of nodules with Acanthoclymenia and Tornoceras that lies 0.1 m (4 in)-thick fossiliferous limestone with auloporid within black shale, 0.38 m (1.25 ft) below the Genundewa. corals, brachiopods, and crinoid fragments. The Lodi is cut Locality 22, Barnes Gully.–A long and complete section out in the main creek section but the horizon lies 0.91 m (3 of the lower Genesee is exposed in Barnes Gully, 2.5 mi (0.4 ft) above the first low falls in the section. The Lodi is ex- km) southeast of Cheshire (Ontario County), on the west- posed in a tributary, 4.1m (13.4 ft) above where the tribu- ern side of Canandaigua Lake, above Foster Point. The sec- tary enters from the north. Above the position of the Lodi tion commences above the Windom/Moscow horizon in the main creek section, a 2 m (6.5 ft)-thick inter- Shale-Geneseo Shale contact at the top of a falls at approx- val of olive gray shale and mudrock is overlain by a 0.61 m imately 800 ft (244 m) altitude. The section illustrates the (2 ft)-thick black shale at the top of a low falls. A Ponticeras great eastward thickening of the Geneseo Shale. No goni- was noted in Bed 21/1c at the erosional base of the black atites were collected at this locality. shale. The black shale is Penn Yan Black Shale B, the higher Locality 23*, Seneca Point Creek.–A section in the upper of the pair of black shales above the Lodi in the Genesee Penn Yan Shale is exposed below the first falls in Seneca Valley (Text-fig. 8), and the first thick black shale above the Point Creek above Seneca Point on the western side of Lodi in the Menteth Gully section (Cd-7) at Canandaigua Canandaigua Lake. Bed 23/a, a bed of concretions 7.1 m Lake (de Witt & Colton, 1978: pls 2-3). (23.3 ft) below the base of the Genundewa Limestone, is Above Penn Yan Black Shale B in Abbey Gulf is a 2.7 m the “hiatus-concretion” horizon of Baird (1976). The hori- (9 ft)-thick interval of siltstone and silty shale (a tongue of zon lies below the level of the pool at the base of the falls, Sherburne Siltstone), at the top of which is a 0.53 m (1.75 0.61 m (2 ft) above a pair of black shales, 0.38 m (1.25 ft) ft)-thick black shale taken to be the Renwick Shale, a unit and 76 mm (3 in) thick. Bed 23/a contains abundant and traceable to Canadaigua Lake (Loc. Cd-7) and eastward to well-preserved pyritic and baritic protoconchs and early its type area near Cayuga Lake where it lies between the whorls of Koenenites and Tornoceras and a distinctive con- Sherburne Siltstone and the Ithaca Shale and Sandstone (de odont fauna with Ancyrodella crosbiensis Kralick, 1994, indi- Witt & Colton, 1978: pls 2-3). cating a correlation with the Crosby Sandstone of Torrey et Above the Renwick Shale, poor goniatites occur in Beds al. (1932) to the east and possibly also with the Linden 21/1a and 21/1b below and above a 0.30 m (1 ft)-thick Horizon to the west; however, the Linden Horizon might black shale, within a 2.6 m (8.5 ft)-thick interval of irregu- be lower in the succession. Bed 23/a and the underlying larly bedded silty shales and siltstones (some with barite and black shales were illustrated by de Witt & Colton (1978: pls poor goniatites); these are the informal “Abbey Beds” of G. 2-4) in the Seneca Creek section (Nap-1). The lower and Baird and C. Brett (Baird et al., 2006: 355; Zambito et al., thicker of the black shales could be the “Starkey” black shale 2007: 94, fig. 4). The “Abbey Beds” are capped by a 0.30 (6 of de Witt & Colton (1978: pls 4-5). 22 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

Locality A, Nap-4, Genundewa Point.–On the eastern Crosby Sandstone found approximately 183 m (600 ft) side of Canandaigua Lake, north of Genundewa Point, the downstream of the Kimble Road bridge. Bed Py-11/b is a equivalent of Bed 23/a of Seneca Point Creek is Bed 23/b, loose block of Crosby with goniatites from approximately a 0.23 m (9 in)-thick concretion bed, beneath a prominent 110 m (360 ft) upstream of the bridge; this bed number is 0.20 m (8 in)-thick siltstone, 5.9 m (19.25 ft) below the used for Luther’s collection. Bed Py-11/c is a loose block of base of the Genundewa Limestone. The horizon and cap- calcareous siltstone with goniatite fragments from approxi- ping siltstone were illustrated by de Witt & Colton (1978: mately 183 m (600 ft) upstream of the second of two dirt pl. 4). tracks which cross the creek approximately 366 m (0.23 mi) Locality B, Ph-5, Shuman Cemetery.–In the tributary of above the bridge. Flint Creek 0.8 mi (1.3 km) north of Shuman Cemetery Locality D, Py-11a, Lakeside Country Club.–Blocks of and 2 mi (3.2 km) south of Gorham (Ontario County), the questionable Crosby Sandstone were found in the creek on Genundewa Limestone, cropping out at approximately 936 the northern side of the Lakeside Country Golf Club, 1.7 mi ft (285 m) elevation, has thinned to a single prominent styli- (2.7 km) southwest of Penn Yan (Yates County), and above olinid limestone several centimeters thick, above black styli- Keuka Lake (East Branch). The locality is below the seventh olinid-rich shales. In the upper Penn Yan Shale, 1.2 m (4 ft) green at approximately 800 ft (244 m) elevation and approx- below the Genundewa, Gordon Baird collected a 10 mm (4 imately 622 m (0.39 mi) west of the Bath Road (NY Rte. in)-thick styliolinid bed with numerous baritic Koenenites 17) and near the Penn Yan-Crosby contact as mapped by styliophilus. The horizon is in a covered interval and must Bergin (1964). be dug out to be collected. The bed could represent either Locality E, Py-14, Willow Grove.–The uppermost Penn the Linden Horizon or the Crosby Sandstone. At the top of Yan Shale, the Crosby Sandstone, and lower tongue of the the Genundewa at this locality, Gordon Baird (pers. comm., Ithaca Shale and Sandstone are exposed in a waterfall section 2002) has found pyritic goniatites in a 1-3 cm-thick corro- in Willow Grove Gully that crosses NY Rte. 54, 3.5 mi (5.6 sional lag marking a discontinuity traceable southeastward km) southwest of Penn Yan (Yates County), above Keuka to Sunset Point ravine on Keuka Lake (see below). This sec- Lake (East Branch). The sections begin at approximately tion was illustrated by de Witt & Colton (1978: pl. 4). 780 ft (238 m) elevation, 0.3 mi (0.48 km) east of the NY Locality C, Py-11, Sartwell Ravine.–The upper Penn Rte. 54 bridge. Bed Py-14/1 is the goniatite-rich basal 10 Yan Shale and lower Ithaca Shale and Sandstone are exposed cm (4 in)-thick unit of the 2 m (6.6 ft)-thick Crosby Sand- in Sartwell Ravine in the vicinity of the Kimble Road bridge stone, a calcareous siltstone with reworked concretions; in- crossing, beside the airport, 1 mi (1.6 km) south of Penn cluded is Tornoceras arcuatum. This section was illustrated by Yan (Yates County). The Crosby Sandstone, at or near the de Witt & Colton (1978: pl. 4). Penn Yan-Ithaca contact, is the source of a large collection Locality F, Py-16, Sunset Point Gully.–The upper Penn of Koenenites styliophilus styliophilus made by D. D. Luther Yan Shale, the Crosby Sandstone, and lower tongue of the in 1904 (his locality 3397) found in the NYSM (Kirch- Ithaca Shale and Sandstone are well exposed in Sunset Point gasser, 1982, 1985). The position of the Crosby Sandstone Gully, above Sunset Point, Keuka Lake (East Branch) that in the section is uncertain. Luther’s label with the collection crosses NY Rte. 54, 0.2 mi (0.32 km) south of Sisson Road notes the horizon at 880 ft (268.2 m) elevation, which and approximately 5.8 mi (9.3 km) southwest of Penn Yan would be approximately 200 yd (183 m) above the Kimble (Yates County). Exposures begin in the upper Penn Yan ap- Road crossing. The Crosby was not found in place in a proximately 15 m (49 ft) east of NY Rte. 54. The 20 cm (8 search in 1977 of the several hundred yards (meters) of in- in)-thick Crosby Sandstone caps the first high waterfall and termittent exposures of the Ithaca upstream of the bridge in the Ithaca beds are well exposed above. the tributary paralleling the new NY Rte. 14A. Bergin Koenenites styliophilus styliophilus is common in the go- (1964: 11) found the Penn Yan-Crosby contact to be well niatite fauna at the base of Bed Py-16/1, the Crosby Sand- exposed at an altitude of 860 ft (262.1 m), although on his stone, a level that also yields Tornoceras arcuatum. A fine map the contact was drawn at approximately 870 ft (265 pyritic specimen of Acanthoclymenia sp. was collected by m), which is at approximately the level of the Kimble Road James Kralick from a concretion in the uppermost Penn Yan bridge and within 10 ft (3 m) of Luther’s locality. De Witt Shale 0.30 m (1 ft) below the base of the Crosby Sandstone. & Colton (1978: pl. 4, table 1) placed the base of the Additional goniatites from the Crosby Sandstone were col- Crosby at 811 ft (247.2 m) elevation, 5.8 m (19 ft) above lected from loose blocks on the creek floor approximately the base of the Renwick Shale, or approximately 21 m (70 69 m (226 ft) east of NY Rte. 54. This section was illustrated ft) below Luther’s locality by de Witt & Colton (1978: pl. 4). Bed Py-11/a consists of loose blocks of goniatite-rich Locality G, Ov-3a, Gully north of Plum Point, Seneca HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 23

Lake.–In an unnamed creek immediately north of Plum unnamed points southeast of Fir Tree Point, approximately Point Road, north of Plum Point, Seneca Lake, 1.3 mi (2.1 1.8 mi (2.9 km) southeast of Rock Stream (Yates County). km) northeast of Himrod (Yates County), poor Ponticeras The access road begins 0.1 mi (0.16 km) south of the inter- occur in a dark silty shale above lenses of pyrite equivalent section of the Bates Cross Road and NY Rte. 14. The beds to the Lodi Limestone (Lodi Bed A) at the Geneseo Shale- of the Lodi Limestone (Lodi Beds A, B, C, and D) are ex- Penn Yan Shale contact. This is locality 12B of Baird et al. posed through a 2.9 m (9.5 ft)-thick interval and, higher in (1989). the Penn Yan Shale, Ponticeras occurs in a 0.18 m (7 in)- Locality H, Ov-4, north of Baskin Point, Seneca thick calcareous siltstone with a fauna of large styliolines, Lake.–On the northern side of Baskin Point, Seneca Lake, gastropods, and brachiopods that lies 2.4 m (8 ft) above the 1.4 mi (2.3 km) northeast of Starkey (Yates County), both top of the Lodi and immediately below a 1.2 m (4 ft)-thick the Lodi Limestone (Lodi Bed B; Bed 25a/1) and the over- black shale (locality 16B of Baird et al., 1989). lying massive siltstone (Bed 25a/2) contain goniatites in a se- Locality L, Ov-10, Mill Creek.–The equivalents of the quence similar to that seen to the south at Fir Tree Point upper Penn Yan Shale and Crosby Sandstone occur in a (Loc. 25) and across the lake at Lodi Glen (Loc. 27). tongue of the Ithaca Shale and Sandstone exposed in Mill Locality I, Ov-5, south of Baskin Point, Seneca Lake.–In Creek, below the Upper Lake Road bridge, 1.3 mi (2.1 km) the creek 0.2 mi (0.32 km) south of Baskin Point, Seneca west of Lodi (Seneca County; eastern side of Seneca Lake). Lake, a single specimen of Ponticeras was recovered in one of The bridge foundation overlies the massive 0.69 m (2.25 the upper cycles of the Lodi Limestone, approximately 5.2 ft)-thick Crosby Sandstone, and approximately 17 m (56 ft) m (17 ft) above the Lodi horizon at the Geneseo Shale-Penn of Penn Yan Shale equivalents are well exposed in the creek Yan Shale contact (sample collected by WTK and Gordon below. Baird). Bed Ov-10/1, with a few well preserved Koenenites stylio- Locality J, Wg-2, north side of Fir Tree Point.–In the philus styliophilus, is believed to be the Linden Horizon. It is embankment, creek, and access road (Rock Stream Road) a nodular band with nautiloids, brachiopods, and auloporids on the northern side of Fir Tree Point, Seneca Lake, poor within a silty shale and mudrock interval that overlies an in- goniatites occur in the styliolinid and ostracod-rich black terval of siltstones (some with ripplemarks) that cap a series shale capping the Fir Tree Pyrite Bed. Above the Fir Tree of falls; the horizon is 6.1 m (20 ft) below the base of the Bed is a 0.68 m (2.25 ft)-thick interval of black shale at the Crosby Sandstone. Bed Ov-10/2 at the top of the silty shale top of the Geneseo Shale (Hubbard Quarry Shale) within and mudrock interval is a 0.28 m (11 in)-thick, irregularly which is a nodular horizon (informal “Bergen Beach Bed” of bedded, silty mudrock with brachiopods, tabulates, and bi- Brett & Baird, 1986). Above is the first of four calcareous valves and a goniatite fragment with the growth lines of beds comprising the Lodi Limestone (Lodi Beds A, B, C, ?Chutoceras nundaium (Hall, 1874). The top of the bed is and D) through an interval of 4.3 m (14 ft). The 0.71 m 4.6 m (15 ft) below the base of the Crosby Sandstone. (2.3 ft)-thick main bed of the Lodi Limestone (Lodi bed B) Poorly preserved Koenenites were noted in Bed Ov-10/3 in crops out 1 m (3.3 ft) above the Fir Tree bed. No goniatites the float immediately beneath a prominent 0.18 m (7 in)- were noted in the Lodi interval at this locality. This is thick siltstone, the top of which is 3.2 m (10.5 ft) below the locality Wg-2 of de Witt & Colton (1978) and locality 16A base of the Crosby Sandstone. Acanthoclymenia occurs in of Baird et al. (1989). Bed Ov-10/4, a silty mudrock, 2.3 m (7.5 ft) below the base Locality 25, Wg-3, Fir Tree Point.–On the western side of the Crosby Sandstone. The horizon lies below a pair of of Seneca Lake, 1.2 mi (1.9 km) southeast of Rock Stream thin siltstones and is 0.91 m (3 ft) below the top of the 0.15 (Yates County), at the southern side of Fir Tree Point (for- m (6 in)-thick ripple-marked siltstone that caps a low falls. merly Rock Stream Landing), the top 1.8 m (6 ft) of the Above the ripple-marked siltstone, Bed Ov-10/5 is a 76 mm Geneseo Shale is exposed (Loc. 25). Above this is a blocky- (3 in)-thick mudrock with a shelly fauna and poorly pre- weathering massive calcareous siltstone approximately 1.8 served Koenenites? that lies 0.91 m (3 ft) below the base of m (6 ft) in thickness (Bed 25/2) with a 0.3 m (1 ft) more the Crosby Sandstone. This section was illustrated by de concretionary unit, below (Bed 25/1) (probably Lodi Lime- Witt & Colton (1978: pl. 3) and Kirchgasser (1981: 45). stone Bed B) and a shale unit above. Locality Wg-3 was il- Locality M, Ov-13, north of Tommy Creek.–Large but lustrated by de Witt & Colton (1978: pl. 5). poorly preserved specimens of Ponticeras cf. perlatum, and Locality K, Wg-7a, Green Point, Seneca Lake.–The rare pyritic goniatite protoconchs in conodont residues, upper Geneseo Shale, Lodi Limestone, and lower Penn Yan occur in a 9 cm (4 in)-thick bed of Lodi Limestone (Lodi Shale are exposed in the bluffs and roadcuts behind cottages Bed C) collected by Gordon Baird at a level 5.5 m (18 ft) along the shore of Seneca Lake between the second and third above the main Lodi Limestone horizon (Lodi B) in the 24 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

gully 0.3 mi (0.48 km) north of Tommy Creek, 2.6 mi (4.2 thors) crops out and the main Lodi Limestone horizon (Lodi km) northwest of Lodi (Seneca County). The main Lodi B) is a fossiliferous calcareous siltstone with auloporid corals Limestone bed (Lodi B) with a pyritic crust at its top, is at near the base of the Penn Yan. Ponticeras perlatum occurs in the top of a waterfall 15 ft (4.6 m) high, and approximately the topmost beds of the Geneseo Shale and in the Lodi bed; 0.30 m (1 ft) above the top of the Geneseo Shale (Hubbard many specimens from both horizons are housed in the PRI Quarry Shale). Pyritic goniatite protoconchs were noted in collections. This section was illustrated by Cole et al. (1959), the conodont residues from the top of Lodi Bed B. De Witt deWitt & Colton (1978: pl. 3), Grasso et al. (1986: fig. 10), & Colton (1978: table 1) placed the top of the Geneseo at Baird & Brett (1986b: 41, fig. 5A; 64, fig. 13), and Thomp- 638 ft (194.5 m) elevation. This is locality 18 of Baird et al. son & Newton (1987: fig. 3). This is locality 22 of Baird et (1989). al. (1989) and Kirchgasser (1994). Locality 27, Lodi Glen.–Above the Tully Limestone in Locality P, Gen-1a, Ovid-Sheldrake Road.–A specimen Lodi Glen (Loc. 3), the Geneseo Shale and Penn Yan Shale of Ponticeras perlatum (CU 41083) was collected by J. W. are exposed in a falls section below the railroad bridge, 1.9 Wells from an outcrop at an elevation of 720 ft (219.5 m) mi (3.0 km) west of Lodi (Seneca County). An interval of on the Ovid-Sheldrake (Parish) Road, approximately 0.7 mi concretions in the lower Penn Yan Shale (or Sherburne Silt- (1.1 km) east of Sheldrake Springs (Seneca County). Wells’s stone) is the type section of the Lodi Limestone, and is the locality 204, below Hall Road, where the creek parallel to source of Ponticeras perlatum, which also occurs, crushed flat, and north of Sheldrake Creek crosses the Ovid-Sheldrake in the shales below. Bed 27/1, the main Lodi bed (Lodi B), Road, is probably in the Lodi Limestone of the lower Sher- with P. perlatum, is a 0.23 m (9 in)-thick concretionary burne Siltstone. The locality was not visited in this study. band, the base of which is 1.2 m (4 ft) above the top of the Locality Q, I-1, Glenwood Creek.–In Glenwood Creek Geneseo Shale (Hubbard Quarry Shale); this is locality 3013 above Glenwood Point, Cayuga Lake, approximately 5 mi (8 of Kirchgasser (1975). Pyritic goniatite protoconchs and km) south of Taughannock Point, Kindle (1896) collected shell fragments are common in the conodont-bone-bed hash specimens referable to Ponticeras perlatum from the upper comprising Lodi Bed A at the contact between the Geneseo Sherburne Siltstone, at his locality 8-1, 12 m (40 ft) below and Penn Yan Shales. A representative of the Epitornoceras the second “Spirifer laevis Zone” (CU 10934b; G125) and peracutum group (E. cf. mithracoides) was collected by J. W. from the second “Spirifer laevis Zone” at the base of the over- Wells at this locality from a level 12.2 m (40 ft) below the lying Renwick Shale, his locality 8-2 (CU 10916; G137- top of the Geneseo Shale. Section Ov-10 was illustrated by 138). de Witt & Colton (1978: pl. 3), Oliver & Klapper (1981b: De Witt & Colton (1978: table 1, pl. 3) recorded the 45, fig. 1), and Grasso et al. (1986: 174, fig. 4). This is lo- base of the Renwick Shale (second “Spirifer laevis Zone” and cality 17 of Baird et al. (1989) and Kirchgasser (1994). Kindle’s locality 8-2) at 634 ft (193.2 m) elevation, which Locality 27a; Ov-16, Romulus Town Quarry.–There are would be upstream of the railroad crossing of the creek. The other sections between Seneca Lake and Cayuga Lake in second “Spirifer laevis Zone” refers to a bed or beds with the which this same interval can be seen. In the large shale pit brachiopod Warrenella laevis (Hall, 1843), which early of the Romulus Town (Ovid) Quarry (Loc. 27a), 1.1 mi (1.8 workers used to define the boundary between the Sherborne km) northwest of Ovid (Seneca County), approximately Siltstone and the Ithaca Shale and Sandstone. Kindle’s Loc. 300 m (984 ft) along a track leading south from West Blaine 8-1 in the underlying Sherburne Siltstone would be at 594 Road, halfway between NY Rte. 96 and the Kinne Road, ft (181.0 m) elevation, above NY Rte. 89 near where the the Lodi Limestone (Lodi B, Bed 27a/1) and associated lev- railroad track crosses the creek. els occur with the typical Ponticeras perlatum fauna. This is The section continues above the Renwick Shale into the locality 3012 of Kirchgasser (1975) and locality 19 of Baird Ithaca Shale and Sandstone. De Witt & Colton (1978) et al. (1989). This section was illustrated by Baird & Brett recorded the top of the Crosby Sandstone at a position 47 (1986b: 48, fig. 9F; 64, fig. 13) and Baird et al. (1989: 364, m (155 ft) above the base of the Renwick at 793 ft (241.7 fig. 7F). m) elevation (downstream of the Duboise Road crossing). Locality 27b; Gen-2, Hubbard Quarry.–The abandoned The section was not examined in this study. shale pit of Hubbard Quarry (Loc. 27b) is immediately east Locality R, I-2, Williams Brook.–A section through the of NY Rte. 98 and just north of Lively Run, 0.12 mi (0.2 Renwick Shale and lower and middle Ithaca Shale and Sand- km) south of the Ovid/Romulus town line and 1 mi (1.6 stone is exposed in Williams Brook, a creek on the south- km) southwest of Kidders (Seneca County). There the upper western side of Cayuga Lake above the site of the old part of the black Geneseo Shale (Hubbard Quarry Shale municipal airport on the northwestern side of Ithaca (Tomp- type section) and lower Penn Yan Shale (Sherburne of au- kins County). Downstream of the NY Rte. 96 bridge is the HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 25

3-m-thick Williams Brook Coquinite of Caster (1933), the above Esty Point at the southeastern end of Cayuga Lake, top of which, at 620 ft (189.0 m) elevation, is approximately approximately 1.3 mi (2.1 km) north of McKinneys (Tomp- 53 m (175 ft) above the base of the Renwick Shale (at 445 kins County), the upper Sherburne is exposed below NY ft or 135.6 m elevation) (de Witt & Colton, 1978: A-16, Rte. 34. The top of the Sherburne indicated by the second table 1). Two specimens in the CU collection referred here “Spirifer laevis Zone” of authors (beds with Warrenella laevis) with question to Chutoceras nundaium might have come has been given as either at 637 ft (194 m) (Williams, 1951) from the Williams Brook Coquinite. The specimens (CU or 616 ft (187.8 m) elevation (Boekenkamp, 1963). From 10928 and 10938) were collected by Kindle (1896) from an unknown level in the Sherburne, F. B. Hine collected (in his locality 4-5, a bed of impure limestone 265 ft (80.7 m) 1876) a Ponticeras perlatum (CU 41079). The section was above the lake. Given a lake level of 382 ft (116.4 m) eleva- not examined in this study. tion, that would place the Williams Brook horizon at 647 ft Locality V, I-11a, Twin Glens at McKinneys.–The upper (197.2 m) elevation, approximately 53 m (175 ft) above the part of the Sherburne Siltstone is exposed in Twin Glens, Renwick Shale, a position close to but higher than that above McKinneys Point at the southeastern end of Cayuga recorded by de Witt & Colton (1978). De Witt & Colton Lake, just north of McKinneys or McKinney’s Station (1978: A-16) placed the Williams Brook Coquinite approx- (Tompkins County). In the northern glen, the second “Spir- imately 21-24 m (70-80 ft) below the Crosby Sandstone. ifer laevis Zone” of authors, which marks the top of the Sher- No new goniatites were found at this locality. burne Siltstone (and base of Renwick Shale), is at Locality S, I-2a, south of Williams Brook.–At a quarry approximately 502 ft (153 m) elevation (Boekenkamp, near the railroad approximately 0.5 mi (0.8 km) south of 1963). Kindle (1896) collected specimens of Ponticeras per- Williams Brook, Kindle (1896) collected a goniatite frag- latum (CU 19030a-b) from his locality 7-4, from the first S. ment here assigned with question to Chutoceras nundaium laevis horizon of the Sherburne, which is approximately 33.5 based on its growth lines and flattened venter. The specimen m (110 ft) below the top of the Sherburne and 20 ft (6.1 m) came from Kindle’s locality 4-2, 4.6 m (15 ft) above the sec- above the lake (approximately 392 ft or 119 m elevation). ond “Spirifer laevis Zone,” a bed or beds with the brachio- From approximately the same horizon (394 ft or 120 m el- pod Warrenella laevis. The horizon is at or near the base of evation), Boekenkamp (1963) collected a Ponticeras (CU the black Renwick Shale. 41075) that is somewhat more involute than typical P. per- Kindle’s locality 4-2 in the vinicity of the base of the latum. These localities were not examined in this study. Gor- Renwick Shale must be near H. S. Williams’ (1884) Station don Baird and Carl Brett have identified a Lodi Limestone 4 locality “at the southwest corner of Cayuga Lake” and pre- horizon with abundant auloporid corals in the basal 3 m (10 sumably near the Spirifer laevis beds at the top of the Sher- ft) of this section at approximately 3 m (10 ft) above lake burne Siltstone and base of the Renwick Shale. Williams’ level (Gordon Baird, pers. comm., 2002). Station 4 yielded specimens referable to Ponticeras perlatum Locality W, I-11c, south of McKinneys.–From a level in (CU 41078-HSW coll. 4A, 41075-HSW Coll. 4B) and an the Sherburne Siltstone at the side of the railroad beginning involute specimen (HSW 4-D35; Cornell locality label, G- approximately 0.5 mi (0.80) south of McKinneys (Tomp- 62) that might be a Koenenites. No new goniatites were kins County) at the southeastern end of Cayuga Lake, Kin- found at this locality. dle (1896) collected specimens of Ponticeras perlatum: CU Locality T, I-11d, Gulf Creek.–From the upper Sher- 10939b and c from his locality 7-2 and CU 10933a and b burne Siltstone in Gulf Creek (Shurger or Shurger’s Glen) from “a short distance to the south” at his locality 7-3. The above Portland Point, approximately 1 mi (1.6 km) south- faunas at Kindle’s localities 7-2 and 7-3 include the aulo- east of South Lansing (Tompkins County), R. P. Boeken- porid coral Cladochonus and could be the faunal horizon of kamp collected a poor specimen of Ponticeras perlatum (CU H. S. William’s (1884) station 48 locality along the side of 41074) at an elevation of 856 ft (260.9 m). The horizon the railroad “a little north of McKinney’s station.” If so, Kin- would be near the NY Rte. 34 crossing of the creek. On a dle’s (1896) localities would be 15 m (50 ft) below the top field map produced by J. W. Wells, the top of the Sherburne of the Sherburne (and base of Renwick Shale) (Williams, Siltstone (second “Spirifer laevis Zone” of authors) is at ap- 1884: 60) or they could be lower horizons, as much as 30 m proximately 940 ft (286 m) elevation, just upstream of the (100 ft) below the top of the Sherburne. In the section above North Triphammer Road crossing, which would place the Willow Point (Loc. I-11), which is approximately 0.5 mi horizon of CU 41074 approximately 26 m (84 ft) below the (0.80 km) south of McKinneys, de Witt & Colton (1978: top of the Sherburne. The section was not examined in this table 1) recorded the top of the Sherburne and base of the study. Renwick at 505.2 ft (155.0 m) elevation. These localities Locality U, I-11b, Esty Glen.–In Esty or Esty’s Glen, were not examined in this study. 26 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

Locality X, Dy-10, Fall Creek, Ithaca.–In the classic sec- in the old Baker Quarry on the Cornell University Campus tion at Fall Creek (Hall, 1843: 250), on the northeastern that was located just above West Road at approximately 710 side of Ithaca (Tompkins County), beginning at the base of ft (216 m) elevation, a specimen presumably collected by J. Ithaca Falls, the Renwick Shale and much of the overlying W. Wells (CU 40106 from his locality 42) is probably a Koe- Ithaca Shale and Sandstone are exposed in a spectacular nenites, as is an uncataloged specimen in the CU collected by gorge. A 0.30 m (1 ft)-thick siltstone cropping out at the S. M. Herrick (SMH-1). The Baker Quarry horizon would level of the pool at the base of the falls marks the top of the appear to correlate with a level within the Triphammer Falls Sherburne Siltstone and from the Warrenella laevis beds in in the Fall Creek section. the lower Renwick Shale approximately 1 m (3 ft) above, Locality Z, Dy-10b, Cascadilla Creek, Ithaca.–In Cas- H. S. Williams (1884: locality 1-B) collected a specimen re- cadilla Creek on the eastern side of Ithaca (Tompkins ferred to Ponticeras perlatum (CU 42289). No new material County), a section of Ithaca Shale and Sandstone is exposed was found at this level here or in the lower Renwick beds in the gorge upstream of the College Avenue bridge. Kindle’s exposed in the Stewart Avenue access road to NY Rte. 13 (1896) locality 2-1 at the base of the lowest cascade in the approximately 0.7 mi (1.1 km) to the north (locality illus- gorge is in dark Renwick Shale. From his locality 2-4, in the trated by Grasso et al., 1986: fig. 16). lower Ithaca beds, 17 m (55 ft) above his locality 2-1, Kindle De Witt & Colton (1978) recognized 13 m (42 ft) of collected a specimen (CU 10932) that might be Chutoceras Renwick Shale in the section but the contact with the over- nundaium. Specimens collected by Kindle (1896) from lying Ithaca beds is inacessable in the falls. Just above the higher in the section (CU 10918; Kindle’s locality 2-10) at crest of the falls on the southern side of the creek is the site the foot of the falls below the College Avenue bridge of the Old University Quarry from which S. G. Williams (Heustis Street bridge and approximately 70 m (225 ft) collected the specimen (NYSM 3753) referred here to above Kindle’s locality 2-1) and NYSM/WTK 3922/1, 2 Chutoceras nundaium. The locality is Williams’ (1884: sec- and CU 10923a-b from 18 m (60 ft) higher (Kindle’s local- tion III, station 3), approximately 72 m (235 ft) above the ity 2-13) are indeterminate. “Spirifer laevis Zone” at the base of the Renwick Shale. The Locality AA, Dy-10c, South Hill, Ithaca.–Two speci- lithology is the distinctive shelly “Firestone bed” facies mens assigned here to Chutoceras nundaium were described (Williams, 1884: 17). In the creek section the horizon proj- by Hall (1879) from a locality in the Ithaca Shale and Sand- ects to the lower part of the “Quarry Sandstone” interval stone at South Hill, Ithaca (Tompkins County). NYSM (Kirchgasser, 1985) that forms the cascade behind the old 3749 and 3751 (the lectotype) were from the Lower electric power plant (Kirchgasser, 1985: 230-231, fig. 5). Chemung Group “at the inclined plane of the railroad” at or A few meters above the top of the power-plant cascade, near Ithaca, New York. The lithology of the enclosing rock a prominent 0.61 m (2 ft)-thick siltstone (“Bluestone Bed” is the shelly Firestone type (Williams, 1884: 17) that yielded of Kirchgasser, 1985) marks the top of the “Quarry Sand- the University Quarry specimen at the edge of Fall Creek, stone” interval. In the first 0.30 m (1 ft) of dark shales im- Ithaca (Loc. Dy-10). The locality could be the same as Kin- mediately below the siltstone bed, Kindle (1896) collected dle’s (1896: 24) locality 5-2, “Quarry at Inclined plane,” goniatites from his locality I-6, one of which is a fine spec- which is 35 m (115 ft) above the Cayuga Street crossing of imen of Koenenites styliophilus (CU 10937; G115). Addi- Sixmile Creek at the base of South Hill. The abandoned tional specimens referred here to K. styliophilus (WTK quarry at approximately 500 ft (152 m) elevation is located 3929/1) have been collected from the 0.30 m (1 ft) of dark below the railroad line and above Stone Quarry Road, just shales beneath the siltstone. The horizon, 82 m (270 ft) to the west of a small northwestward-flowing creek. No new above the base of the Renwick, is believed to be the Linden goniatites were found at the locality. Horizon of the upper Penn Yan Shale in western sections Locality BB, I-8a, Enfield Glen.–An uncataloged speci- (Kirchgasser, 1985). Others specimens of K. styliophilus from men in the CU collection from above the top of the lower higher in the section are NYSM 12165/WTK 3931/1, a falls in Enfield Glen, approximately 4 mi (6.4 km) south- specimen found in float on top of the siltstone ledge, and a west of Ithaca (Tompkins County), is assigned here with specimen (CU 10925) collected by Kindle (1896) from his question to Chutoceras nundaium. The specimen collected locality I-7 in the floor of the creek above the siltstone. Ad- by K. E. Caster or G. Q. Williams came from within the ditional but poor specimens of Koenenites include CU Ithaca Shale and Sandstone given that the base of the 13+m 10929 from Kindle’s locality I-8, just below the base of (44+ft) Renwick Shale at approximately 448 ft (136 m) el- Triphammer Falls. The upper part of section was illustrated evation in the Cayuga Inlet (Loc. I-8, Lick Brook: de Witt by Kirchgasser (1985: fig. 5; see Addendum). & Colton, 1978: table 1) is at least 50 ft (152 m) below Locality Y, Dy-10a, Baker Quarry.–From the Ithaca beds where the section begins near the base of the first falls in the HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 27

gorge. No attempt was made here to locate the goniatite County, the nodular beds of lower Genundewa are progres- horizon. sively cut out leaving only a single well-bedded upper Ge- Locality CC, Gen 13a, Big Salmon Creek, near nundewa horizon exposed in Erie County. Genoa.–A Tornoceras? was noted in the pyrite capping the The nodular beds of the Genundewa mark the entry of Fir Tree Limestone of the upper Geneseo Shale in the north- Tornoceras uniangulare compressum Clarke, 1898, Koenenites ern of two closely spaced westward-flowing tributaries of Big styliophilus kilfoylei n. ssp., and Acanthoclymenia genundewa Salmon Creek, southwest of Genoa (Cayuga County). Sam- (Clarke, 1898). Also there is K.? fasciculatus (Clarke, 1898), ple WTK 3978 was given by Gordon Baird. This is locality which is based on a single specimen collected at Genundewa 32 of Baird et al. (1989: fig. 4F, horizon d) and Kirchgasser Point, Canandiagua Lake (Loc. 23b). The Manticoceras (1994). lamed group (sensu House & Ziegler, 1977), in the form of Locality DD, Gen-14a, Big Salmon Creek, Venice Cen- M. contractum (Clarke, 1898), is now believed to enter the ter.–A goniatite fragment and protoconch of Ponticeras? was succession above the nodular beds, in the more regularly found in the pyrite capping the Fir Tree Limestone exposed bedded upper Genundewa. M. sinuosum apprimatum near the mouth of the southern fork of an unnamed west- Clarke, 1898, which is similar to M. contractum, might also ward-flowing tributary of Big Salmon Creek, northeast of prove to enter in the topmost bed(s) of the Genundewa. The Venice Center (Cayuga County). The sample (WTK 3989) position in the Genundewa of M. nodifer (Clarke, 1885) is was provided to us by G. Baird from locality 31 of Baird et unknown. al. (1989: section 4E, horizon d) and Kirchgasser (1994). Because of the thinness of the Genundewa interval, ex- Locality 28. Pleasant Valley, near Sherburne.–The Gene- panded scales on Text-fig. 8 are used in sections west of seo Shale crops out in the creek immediately opposite Pleas- White Creek (Loc. 16); these are indicated by black dots ant Valley Bible Baptist Church, in the creek just south of and open circles next to the solid vertical band beside the Pleasant Valley Road, 1.9 mi (3.0 km) south-southeast of sections. Surfaces of submarine erosion associated with the Sherburne (Chenango County). From here, J. W. Wells Genundewa Limestone in the condensed Genesee sections made a good collection of tornoceratids and ponticeratids, near the basin margin in Erie County have been docu- and one specimen of Pharciceras? sp. (NYSM 12128; see Pl. mented in detail by Brett & Baird (1982, 1990, 1996, 1997) 32, Fig. 14) is known. Gordon Baird (pers. comm., 2002) and Baird & Brett (1986a, b) (Text-fig. 7). The sub-Genun- recently visited this locality and recognized a few feet of the dewa disconformity with the underlying North Evans Lime- Fillmore Glen Member of the Tully Limestone at the base of stone (Conodont bed of Hinde, 1879), which is at the base the section (probable source of the Pharciceras? sp.), followed of the upper Genundewa in Erie County, serves to separate by approximately 3 m (12 ft) of Geneseo Shale, which is the nodular lower Genundewa from the more regularly bed- abruptly succeeded by massive beds of Sherburne Siltstone ded upper Genundewa farther east between Bethany Center exposed upstream of the church area. (Loc. 24), Linden (Loc. 15), and Linden Falls (Loc. 15a) in Genesee County, and Fall Brook (Loc. 19) in the Genesee Genundewa Limestone Valley in Livingston County. Between the Genesee Valley Although relatively thin, the Genundewa Limestone (Styli- and the type section at Canandaigua Lake, the shale in- ola Limestone of early authors) is the only widespread lime- terbeds in the Genundewa interval thicken and the differen- stone unit above the Tully in New York and the oldest tiation of the Genundewa into lower and upper parts is less horizon with a rich and diverse goniatite fauna. Unfortu- apparent. nately, because of generally poor preservation due to recrys- East of Canandaigua Lake where the Penn Yan Shale and tallization and sporadic occurrence, the fauna is still West River Shale pass into the slope facies of the Ithaca Shale imperfectly known. At the type locality, at Genundewa and Sandstone, the Genundewa Limestone horizon is diffi- Point on the eastern side of Canandaigua Lake, de Witt & cult to locate. At Shuman Cemetery, approximately midway Colton (1959) recognized 3.7 m (12 ft) of Genundewa between Canandaigua Lake and Keuka Lake, the Genun- Limestone consisting of several 25-250 mm (1-10 in)-thick dewa is represented by a single styliolinid bed several cm nodular styliolinid layers separated by dark gray shale. West- thick (locality Ph-5 of de Witt & Colton, 1978: pl. 4) and ward toward Lake Erie, with the thinning and eventual loss capped by a 1 in (25 mm)-thick lag bed. This styliolinid bed of the shale beds, the nodular beds converge to form an ir- was thought by de Witt & Colton (1978) and earlier work- regularly bedded limestone (informal lower Genundewa ers to be the easternmost occurrence of the unit. From Limestone) overlain by a single well-bedded limestone (in- Keuka Lake southeastward to Cayuga Lake, de Witt & formal upper Genundewa Limestone often exposed as the Colton (1959, 1978: pls 3-4) correlated the Genundewa caprock of waterfalls. West of Linden (Loc. 15), in Genesee with the Crosby Sandstone of Fox (1932) and Torrey et al. 28 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

(1932), and used the base of the Crosby as a datum within presumed North Evans Limestone (Kirchgasser, 2002), a the thick and undifferentiated silty faces of the Ithaca Shale level marking the hiatus between the 23-cm-thick nodular and Sandstone. As noted earlier (Kirchgasser, 1985), Gor- lower Genundewa and the 18-cm-thick, more regularly bed- don Baird discovered the Genundewa horizon at Keuka ded, upper Genundewa. Lake in Ithaca facies 5-9 m (16-30 ft) above the Crosby. At Locality 15a, Linden Falls.–At the main falls in the Little Penn Yan (DDs; Py-10/1), Willow Grove (EE; Py-14/2), Tonawanda Creek at Linden (Genesee County), the Genun- and Sunset Point (FF; Py-16/2), the bed is a pyritic, nodular dewa Limestone is 0.36 m (14 in) thick. Goniatites are discontinuity horizon with goniatites and conodonts that abundant in Bed 15a/12, the lowermost 76 mm (3 in) of are restricted to the lower and upper Genundewa farther the 0.18 m (7 in)-thick nodular lower Genundewa and in west. the overlying Bed 15a/13, a 76 mm (3 in)-thick bed in the Still farther southeast at Seneca Lake, in the creek east of middle of the lower Genundewa interval. Only a few goni- Starkey (Yates County), a 5-cm-thick, cross-laminated and atites were noted in the more uniformly bedded and resist- pyritic styliolinid bed identified by Gordon Baird as Genun- ant 0.18 m (7 in)-thick interval of the upper Genundewa dewa lies within Ithaca facies 16 m (53 ft) above the base of that caps the falls. the Crosby. No goniatites have been found in the bed, which Locality 16*, White Creek.–A 0.34 m (1.1 ft)-thick sec- crops out in the creek approximately 30 yd (27 m) south of tion of Genundewa Limestone crops out in the bed of the NY Rte. 14 (locality GG; OV-6 of de Witt & Colton, western branch of White Creek in a field north of NY Rte. 1978). It is unclear whether the horizons identified as Ge- 20, 2.1 mi (3.4 km) east of Bethany Center (Genesee nundewa at Keuka Lake and Seneca Lake represent a single County). A few poor specimens of Koenenites were noted in condensed record of the entire Genundewa interval to the Bed 16/8, the 0.08-0.09 m (3-4 in)-thick lower nodular in- west or only one of possibly several cryptic Genundewa hori- terval of the Genundewa. The upper, more regularly bedded zons that splay into the southeastward-thickening Ithaca interval is 0.26 m (10 in) thick and poorly preserved baritic Shale and Sandstone; for discussion of the genesis of the goniatites occur in Bed 16/9, the uppermost 76 mm (3 in) somewhat similar Fir Tree and Lodi Limestone horizons, see of the unit. Baird et al. (1989). The position of the Genundewa and its Locality 17*, Taunton Gully.–At the top of the upper fauna is unknown in the Ithaca Shale and Sandstone at falls in Taunton Gully, 1.7 mi (2.7 km) north of Leicester Cayuga Lake but the interval probably projects to a level (Livingston County), below the bridge where Starr Road high in that unit (Kirchgasser, 1985: fig. 3). crosses the creek, the Genundewa Limestone is 0.56 m (1 ft 10 in) thick. Above a 0.13 m (5 in)-thick black shale that Localities.–Locality numbers with asterisks have sections il- marks the top of the Penn Yan Shale, the lowermost 0.25 m lustrated in Text-figs 8-9. (10 in) of the Genundewa consists of nodular limestone. Locality 13*, Cayuga Creek.–A poorly preserved Koe- Bed 17/8 at the top of this nodular interval is the level that nenites was noted in the topmost 25 mm (1 in) shaley layer produces Acanthoclymenia genundewa and Tornoceras. The (Bed 13/9) of the 0.2 m (8 in)-thick Genundewa Limestone overlying 0.25 m (10 in) of more uniformly bedded upper at the top of the falls over the Genundewa in Cayuga Creek, Genundewa is sample-horizon 4586-SD of de Witt & 2 mi (3.2 km) northwest of Cowlesville (Wyoming County). Colton (1978) and Huddle (1981: B53). Locality 14a, Murder Creek.–In Murder Creek at Gris- Locality 18*, Beards Creek.–The Genundewa Limestone wold (Genesee County), 0.8 mi (1.3 km) south of Darien, caps the spectacular falls in Beards Creek, approximately 175 goniatites are abundant in the basal bed, 51-76 mm (2-3 in) yd (158 m), downstream from the Dunkley Road bridge, thick, of the Genundewa Limestone (Bed 14a/3), which 0.8 mi (1.3 km) northwest of Leicester (Livingston County). crops out at the level of a farm road that crosses the stream The nodules of Bed 18/6a produce Acanthoclymenia genun- 34 m (37 yd) upstream of the culvert beneath the railroad dewa, rare Koenenites styliophilus kilfoylei n. ssp., and tracks. This locality was noted by Clarke (1899c: 108-109). Tornoceras; the bed lies 0.15 m (6 in) above the lip of the Locality 15*, Linden.–The Genundewa Limestone falls, at the top of the 0.15 m (6 in)-thick nodular lower Ge- forms the caprock of the waterfall in the small eastward- nundewa Limestone; this is locality 3001 of Kirchgasser flowing tributary to Little Tonawanda Creek, 0.3 mi (0.48 (1975: 68). The overlying 0.25 m (10 in)-thick upper Ge- km) north of the bridge at Linden (Genesee County). In the nundewa is more massive and regularly bedded. 33-cm-thick exposure a few yards (meters) south of the falls, Locality 19, Fall Brook-Dewey Hill.–At the top of the Koenenites and Tornoceras are common in the 10 cm nodular NY Rte. 39-20A roadcut at Dewey Hill, 1 mi (1.6 km) bed at the base of the unit. At the top of the overlying 5 cm south of Geneseo (Livingston County), the lowermost 0.25 nodular bed, pyritic bactritids and protoconchs and early m (10 in) of nodular Genundewa Limestone is exposed on whorls of Koenenites occur in microfossil residues from the HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 29

the southern side of the road above a 0.13 m (5 in)-thick falls, 1.25 m (4.1 ft) above a 0.13 m (5 in)-thick black shale black shale overlying a prominent 0.1 m (4 in)-thick, well- that marks the top of the Penn Yan Shale, and 1.17 m (3.8 bedded argillaceous limestone at the top of the Penn Yan. ft) below the highest styliolinid bed; this is locality 3006a of Poor Koenenites and Acanthoclymenia occur in the nodules of Kirchgasser (1975: 68). The fauna of Bed 23/1 includes Bed 19/10a at the base of the interval; this horizon is bed Acanthoclymenia, Koenenites, and Tornoceras. Tornoceras is DH-1 of Kralick (1991, 1994) and is approximately the particularly abundant in the correlative of Bed 23/1 in level of sample-horizon 8962-SD of Huddle (1981: B52). Miller’s Creek, Canandaigua Lake, 0.5 mi (0.8 km) south Bed 19/10, at the top of the nodular interval, is a 0.05 m (2 of Barnes Gully (Bed MC-3 of Kralick, 1991: fig. 2; 1994: in)-thick nodule bed with fragments of Manticoceras con- 1395). tractum?, Acanthoclymenia, and Tornoceras. Bed 19/10 corre- Locality 23b, Genundewa Point, Genundewa sponds to bed DH-2 of Kralick (1991, 1994) and Lake.–This section is along the eastern side of Canandaigua sample-horizon 6681-SD of de Witt & Colton (1978) and Lake, north of Genundewa Point (cottage number 900), Huddle (1981: B52). below East Lake Road, 1 mi (1.6 km) north of Vine Valley In the side creek at Fall Brook, adjacent to Dewey Hill, (Yates County). Tornoceras sp., Acanthocylmenia genundewa, the complete Genundewa Limestone is approximately 0.71 and Koenenites? fasciculatus occur along with abundant bac- m (2.3 ft) thick. In the accessible section along the footpath tritids in the styliolinid lenses of Bed 23b/4 cropping out 100 yd (91 m) north of the main falls, Bed 19/10 with Acan- between 0.71 m (2.3 ft) and 0.84 m (2.75 ft) above the 0.1 thoclymenia genundewa is a 0.12 m (5 in)-thick nodular bed m (4 in)-thick black shale that defines the base of the 2.4 m at the top of the lower nodular interval of the Genundewa (8 ft)-thick Genundewa interval. Tornoceras is relatively in the recess beneath the well-bedded 0.17 m (7 in)-thick common in the fossiliferous and baritic styliolinid ledge upper Genundewa; here the base of Bed 19/10 is 0.32 m (1 (Bed 23b/5) that lies between 1.14 and 1.45 m (3.75 and ft) above the base of the Genundewa, and 0.29 m (1 ft) 4.75 ft) above the base of the black shale at the base of the below the top of the Genundewa Limestone as here defined; Genundewa; this is sample-horizon 6718-SD of de Witt & this is bed FB-1 of Kralick (1991, 1994). Colton (1978: Section Nap-4, pl. 4) and Huddle (1981: Locality 19a, Fall Brook, loose blocks.–Loose blocks of B51) and the correlative of Bed 23/1 in the Seneca Point Genundewa Limestone occur at the base of the high falls in section. Poor baritic goniatites were noted in the uppermost Fall Brook, 1.5 mi (2.4 km) south of Geneseo (Livingston styliolinid ledge (Bed 23b/6), which dips below lake level County). The blocks contain baritic specimens assigned to approximately 50 yd (46 m) north of the point. Bed 23b/6 Manticoceras contractum and are believed to have come from (=3958-SD) lies between 1.96 and 2.13 m (6.4 and 7 ft) the topmost beds of the Genundewa. above the base of the Genundewa; the top of Bed 23b/6 im- Locality 22, Barnes Gully.–In Barnes Gully, above Foster mediately underlies a 0.3 m (1 ft)-thick interval of thin styli- Point on the western side of Canandaigua Lake, 2.5 mi (4.0 olinid lenses that is taken as the top of the Genundewa km) southeast of Cheshire (Ontario County), the Genun- Limestone. dewa Limestone consists of a sequence of styliolinid lime- Locality 24, Bethany Center.–Numerous loose blocks of stone horizons. Bed 22/7, 1.14 m (3.75 ft) below the Genundewa Limestone cover the embankment below NY topmost of the styliolinid units and 1.2 m (4 ft) above a 0.3 Rte. 20 for several hundred yards (meters) east of the Center m (1 ft)-thick black shale, has barite and poorly preserved Road bridge that crosses NY Rte. 20 at Bethany Center goniatites. Here the black shale band is taken to mark the (Genesee County). The goniatites Koenenites styliophilus kil- base of the Genundewa because this is the horizon that can foylei n. ssp. and Tornoceras uniangulare compressum are be seen below the Genundewa to the west. There are styli- abundant in certain levels, but unfortunately most of the olinid limestone horizons lower than this, both here and, blocks are highly weathered and the already recrystallized for example, at Seneca Point Creek. The top of the Genun- specimens tend to crumble. The most productive horizon is dewa is at 946 ft (288.3 m) altitude (de Witt & Colton, Bed 24b/1 of the Genundewa outcrop below the Center 1978: table 1). Road overpass (Loc. 24b). Small pyritic specimens of Acan- Locality 23, Seneca Point Creek.–In Seneca Point Creek, thoclymenia occur in the shaley styliolinid beds (Bed 24/2) above Seneca Point on the western side of Canandaigua Lake on the tops of loose blocks on the southern side of the high- (Ontario County), the styliolinid limestones that comprise way (including specimens donated by James Kradyna). The the Genundewa Limestone extend through an interval of horizon is probably from the topmost 76 mm (3 in) of the 2.4 m (7.9 ft) and are exposed in the face of the first falls Genundewa exposed beneath the bridge. above the lake (Text-fig. 9). Bed 23/1 is a 0.23 m (9 in)- Locality 24b, Bethany Center roadcut.–The upper beds thick styliolinid unit with barite, which forms the lip of the of the Penn Yan and the overlying Genundewa Limestone 30 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

are seen in the roadcut below the Center Road overpass of stone that crops out in the creek 2 m (7 ft) below a massive NY Rte. 20 at Bethany Center (Genesee County). The Ge- siltstone and approximately 30 yd (27 m) south of NY Rte. nundewa exposure is directly beneath the bridge, and here 14. De Witt & Colton (1978: table 1, pl. 5) recognized 0.46 the limestone is 0.38-0.40 m (1.25 ft) thick. Goniatites are m (1.5 ft) of Crosby Sandstone at this locality with the base most abundant in the 0.12-0.15 m (5-6 in)-thick nodular cropping out at 577.5 ft (176.0 m) elevation. No goniatites lower Genundewa (Bed 24b/1). were found in the Genundewa at this locality. Locality DD, Py-10, Cornwall Gully.–In Cornwall Gully, above NY Rte. 364 on the northwestern side of Penn West River Shale Yan (Yates County), the Genundewa Limestone is repre- This unit comprises the sequence of medium gray shales and sented by a pyritic lag deposit (Bed Py-10/1) within a mudrock with numerous black bands that lies between the tongue of the Ithaca Shale and Sandstone. The horizon is a Genundewa Limestone and the black Middlesex Shale. Go- thin band of rusty pyritic nodules with poorly preserved niatites occur both as pyritic internal molds and molds Koenenites? fasciculatus and Acanthoclymenia within a 0.79 within the common concretions, approximately 0.3 m (1 ft) m (2.6 ft)-thick shaly interval, which crops out on the east- in diameter, of which there are several levels in the sequence. ern side of the creek above a series of low falls over siltstones, Goniatites also commonly occur crushed flat within the between 5.2 and 6.1 m (17 and 20 ft) above the base of the shale. The unit is 2.4 m (8 ft) thick on Lake Erie near Pike Crosby Sandstone. The 2 m (7 ft)-thick Crosby Sandstone Creek outlet; on the western side of the Genesee Valley it is at the base of the tongue of the Ithaca Shale and Sandstone 16.2 m (53 ft) thick and at Seneca Point Creek approxi- crops out at approximately 860 ft (262 m) elevation (de mately 40 m (132 ft) thick. East of Canandaigua Lake, the Witt & Colton, 1978: table 1, pl. 4). This locality was found West River Shale interfingers with the upper part of the by G. Baird. Ithaca Shale and Sandstone and partly overlies that unit. Locality EE, Py-14, Willow Grove Gully.–In Willow The West River Shale is more extensive than as originally Grove Gully, Keuka Lake, above NY Rte. 54, 3.5 mi (5.6 defined by Clarke & Luther (1904); the current definition km) southwest of Penn Yan (Yates County), the horizon of is that of de Witt & Colton (1959: 2819). Full details of the Genundewa Limestone is represented by Bed Py-14/2, thickness and lithological variation were given by de Witt a thin concretionary bed, with pyritic seams and cross sec- & Colton (1978). Meter-scale cycles of alternating black tions of Koenenites, that lies within a tongue of the Ithaca and gray shale with concretions characterize the West River Shale and Sandstone, approximately 8.8 m (29 ft) above the Shale particularly between the Genesee Valley and Keuka base of the Crosby Sandstone. The horizon is 0.22 m (9 in) Lake. On the correlation charts of de Witt & Colton (1978: below a prominent 0.34 m (1.2 ft)-thick black shale; the pls 2, 4), 23 cycles can be recognized from no. 1 above the black shale is the first black shale above the Crosby Sand- top of the Genundewa Limestone to no. 23 immediately stone illustrated by de Witt & Colton (1978: pl. 4). below the base of the Middlesex Shale. Locality FF, Py-16, Sunset Point Gully.–In Sunset Point A useful lithic marker in the upper West River Member Gully, Keuka Lake, above NY Rte. 54, 5.8 mi (9.3 km) is the Bluff Point Siltstone Bed (Torrey et al., 1932; Bergin, southeast of Penn Yan (Yates County), the horizon of the 1964; de Witt & Colton, 1978). This horizon, with distinc- Genundewa Limestone is represented by a thin concre- tive convolute bedding, ranges in thickness from 1.5 to 11 tionary bed (Py-16/2) with pyritic seams within a tongue of cm (0.05 ft to 0.35 ft) (de Witt & Colton, 1978: table 1). the Ithaca Shale and Sandstone, approximately 8.5 m (28 It lies within the fourteenth cycle and can be seen in most ft) above the base of the Crosby Sandstone. Small pyritic go- sections between the Genesee Valley and Seneca Lake. niatites are common in the bed and include the typical Ge- The following sequence of goniatities is recognized in nundewa forms Koenenites? fasciculatus, Acanthoclymenia cf. the area between the Genesee Valley and Keuka Lake. Koe- genundewa, and Tornoceras. Bed Py-16/2 is 0.91 m (3 ft) nenites sp., Acanthoclymenia sp., and Tornoceras sp. occur below a 0.48 m (1.6 ft)-thick black shale that is the first sporadically through the lower third of the member. The black shale above the Crosby Sandstone illustrated by de fauna of the middle part of the member up to and including Witt & Colton (1978: pl. 4). the relatively fossiliferous shales surrounding the concre- Locality GG, Ov-6, creek east of Starkey.–In the creek tionary bed, in the thirteenth cycle, just below the thin dis- east of Starkey (Yates County), and south of Brewer Point, tinctive convolute-bedded Bluff Point Siltstone Bed (Bergin, Seneca Lake, G. Baird located the Genundewa Limestone 1964; de Witt & Colton, 1978), is dominated by Koenenites equivalent at a position 16 m (53 ft) above the base of the beckeri n. sp. along with K.? cf. fasciculatus, and Loboto- 1.4 m (4.6 ft)-thick Crosby Sandstone. The horizon is a 5 rnoceras aff. hassoni House, 1978. cm (2 in)-thick cross-laminated, pyritic, styliolinid lime- The uppermost West River Member contains a distinct HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 31

goniatite fauna, particularly in the septarian horizons such 1.5 m (5 ft) across, 2.4 m (8 ft) below the base of the Mid- as those 2.4 and 3.4 m (8 and 11 ft) below the Middlesex dlesex Shale. Koenenites beckeri n. sp. form D and Ponticeras? Shale in Beards Creek (Loc. 18a/4b in cycle 18, the sp. occur in Bed 18a/5. Williamsburgh Bed of Over et al., 1999, 2003, and 18a/5 in Locality 19a*, Fall Brook.–The Fall Brook section con- cycle 19–20, the Beards Creek Horizon of Kirchgasser & tinues upward from that shown in Text-fig. 8 (Loc. 19). Bed House, 1981). Here are found K. beckeri n. sp. form D, 19a/1 is the shale unit 0.9 m (3 ft) above the Genundewa Manticoceras lamed aff. cordatum (G. & F. Sandberger, immediately above the falls. Beds 19a/2-4 are above the 1850), Acanthoclymenia aff. neapolitana (Clarke, 1892), highway bridge of NY Rte. 63. Bed 19a/2 is approximately Acanthoclymenia sp. A (Kirchgasser, 1975; Loc. 26a/1), Pon- 0.5 m (2 ft) upward in the exposure above the bridge, per- ticeras? sp., K.? cf. fasciculatus, and Tornoceras sp. Bed 18a/4b haps 2.3 m (7.5 ft) above the Genundewa, and the goniatites at Beards Creek, named the Williamburgh Bed by Over et are crushed in a capping seam of a thin Styliolina-rich unit; al. (1999: B15; 2003: 219) is a styliolinid grainstone, rich in this is sample-horizon 8122-SD of de Witt & Colton (1978: pyrite, wood, and conodonts, which is intermittently ex- pl. 2) and Huddle (1981: B52) (Kirchgasser et al. 1994: fig. posed near the top of the West River Shale between Lake 10). Bed 19a/3 is from dark shale approximately 4.7 m (15.5 Erie and the Genesee Valley. To the east, goniatites are found ft) higher, and Bed 19a/4 is a large septarian level approxi- in the Williamsburgh Bed at Whetstone Brook (Bed 26a/1), mately 2.0 m (6.5 ft) higher again. near Honeoye (Ontario County); the same horizon might be Locality 23a, Seneca Point Creek.–There is a long sec- Py-7/2 in Chidsey Point Creek, Keuka Lake, in Steuben tion of the West River Shale in Seneca Point Creek, above County. Seneca Point, on the western side of Canandaigua Lake (On- tario County). At the top there is a major falls over the black Localities.–Locality numbers with asterisks have sections il- Middlesex Shale. Bed 23a/3 is a concretionary level at the lustrated in Text-fig. 9. The goniatites from the West River top of cycle 13, which is overlain by a 0.18 m (7 in)-thick Shale are mostly poorly preserved and squashed flat, apart black shale at the base of cycle 14. The horizon crops out from a few horizons with pyritic inner whorls and calcitic or approximately 60 yd (54 m) downstream from the foot of baritic replacements. The distribution of goniatites in this the falls and just above a cascade where a side creek enters unit has not been investigated comprehensively. from the north. Bed 23a/3 is 0.67 m (2.2 ft) below the 1.5 Locality 18a*, Beards Creek.–The Beards Creek section cm (0.6 in) Bluff Point Siltstone in cycle 14 (see de Witt & continues upward from that illustrated in Text-fig. 8 (Loc. Colton, 1978: pls 3-4). Bed 23a/3 yields Koenenites beckeri 18) and the levels that yielded goniatites are indicated in n. sp. and Lobotornoceras aff. hassoni. Lower, another side Text-fig. 9. Beds 18a/1-3 are exposed below the Dunkley creek enters from the north, and Bed 23a/2, a concretion Road bridge. Bed 18a/1 is 0.15 m (6 in) above the Genun- horizon exposed approximately 30 yd (27 m) below this, is dewa, Bed 18a/2, 0.6 m (2 ft) above, and Bed 18a/3 approx- in cycle 13, 2.7 m (8.75 ft) below the Bluff Point Siltstone. imately 2.13 m (7 ft) above, and 0.3 m (1 ft) below the Farther downstream, a prominent 0.38 m (1.25 ft)-thick foundation of the bridge on the eastern side of the road. black band is well exposed in the floor of the creek upstream Above the road, Bed 18a/4 is a line of large septaria nodules of a low falls, and Bed 23a/1 with K. sp. is a concretion bed with goniatites in adjacent shales beneath a 0.17 m (7 in)- in the cycle below (cycle 12), approximately 5.3 m (17.5 ft thick black shale approximately 7 m (23 ft) below the Mid- below the Bluff Point Siltstone. In this section, the Bluff dlesex Shale; the bed is in the cycle below the one that Point Siltstone is approximately 15.5 m (51 ft) below the includes the Bluff Point Siltstone (de Witt & Colton, 1978: base of the Middlesex Shale (de Witt & Colton, 1978). pls 2, 6) but the key marker horizon was not seen in the sec- Locality 26a*, Whetstone Brook.–The upper part of the tion. In the succeeding cycle, which elsewhere includes the West River Shale is exposed in Whetstone Brook down- Bluff Point Siltstone, Bed 18a/4a (cycle 15) is a gray stream from the crossing of Huff Road (which passes west to mudrock with Koenenites beckeri n. sp., 4.9 m (16 ft) below Big Tree Road), 1.6 mi (2.5 km) northwest of Honeoye the base of the Middlesex Shale. Near the top of the section, (Ontario County). The section illustrated here below the Bed 18a/4b (Williamsburgh Bed) in cycle 18 is a Fossil Log Middlesex Shale commences at approximately 1,035 ft (315 bed (concretionized raft of driftwood), 3.4 m (11 ft) below m) altitude (Text-fig. 9). Two horizons consist of a large- the base of the Middlesex Shale which underlies a 0.3 (1 ft) septarian nodule horizon (Bed 26a/2) above, and a smaller black shale; this linguloid-rich bed yields Manticoceras lamed nodule horizon (Bed 26a/1), 0.76 (2.5 ft) below a 0.3 m (1 aff. cordatum, K.? cf. fasciculatus, Ponticeras? sp., Acanthocly- ft)-thick black shale unit; Bed 26a/1, with Acanthoclymenia menia aff. neapolitana, and Tornoceras. Bed 18a/5, the Beards sp. A and Lobotornoceras aff. hassoni, is locality 3003 of Creek Horizon, in cycle 19-20, is a line of vast septaria to Kirchgasser (1975: 68) and the Williamsburgh Bed of Over 32 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

et al. (1999, 2003). The key black shale, at the base of West the Bluff Point Siltstone (Bed Ov-10/6) is a dark gray shale River cycle 18, lies 6.1 m (20 ft) below the base of the Mid- immediately above a 10 in (0.25 m)-thick black shale. The dlesex Shale (de Witt & Colton, 1978: pl. 2) and can be horizon, at approximately 905 ft (276 m) elevation, is 3.6 m traced on the correlation charts of de Witt & Colton (1978: (11.8 ft) below the Bluff Point Siltstone. The black shale pls 2, 4) eastward to Canandaigua Lake and westward to the and overlying Bluff Point Siltstone were illustrated by de Genesee Valley where at Beards Creek (Loc. 18a) it is the Witt & Colton (1978: pl. 3); see also Oliver & Klapper boundary between cycle 18 with the Fossil Log concre- (1981b: 45, fig. 1, sample WTK 3937). tionary bed (18a/4b) and the Beards Creek Horizon (18a/5) Locality HH, Nap-3, Granger Point Road.–The upper in cycle 19-20 (Text-fig. 9). Poor pyritic goniatites occur in West River Shale is well exposed in the roadcut and side the dark shales of Bed 26a/3, approximately 0.91 m (3 ft) creek along NY Rte. 21 across from the road to Granger below the base of the Middlesex Shale. Point, Canandaigua Lake (Ontario County). The Bluff Locality 29a, Hicks Point.–This is the locality on the Point Siltstone is approximately 4 m (13 ft) above a 0.5 m western side of Canandaigua Lake (Ontario County) from (1.6 ft)-thick dark gray to black shale near the base of the which Wells (1956b) reported the first specimen of the section. The Bluff Point Bed is 0.5 m (1.6 ft) above a 0.15 genus Koenenites known in New York. The section at the m (6 in)-thick black shale, and specimens of Koenenites in southwestern side of Hicks Point begins at lake level and dark shales and concretions in the float at the base of the continues in the bluffs below and above Bopple Hill Road. section might have come from the cycle below the 0.15 m The locality was visited by MRH with J. W. Wells in 1959, (6 in)-thick black shale. and, although the specimen was loose, it was thought it Locality II(XX), Nap-8, 0.8 mi south of could have come from the shale interval 5.5-7.3 m (18-24 Middlesex.–This section in the West River Shale is in the ft) above the lake level, and below the two septarian horizons westward-flowing creek that crosses NY Rte. 245, 0.8 mi higher in the cliff. It could also have come from Bed 29a/1, (1.3 km) south of Middlesex (Yates County). Approximately a 0.46 m (1.5 ft)-thick band of dark gray to black silty shale 46 ft (14 m) upstream from where the section commences approximately 2.6 m (8.5 ft) above lake level that yields sim- is a band of small concretions in the creek floor and the 50 ilarly preserved crushed molds of Koenenites and Acanthocly- mm (2 in)-thick convolute-bedded Bluff Point Siltstone lies menia. Bed 29a/1 is within the section of gray mudrock 6.3 m (21 ft) above. Bed Nap-8/1 (WTK 3884) is a fossil- exposed in the shale pit below the road and just west of the log type concretion with pyrite and barite seams with a creek. fauna of styliolinids, gastropods, small lingulid brachiopods, Locality 46c, Snyder’s Gully.–The upper beds of the and goniatites. The loose concretion is believed to have West River Shale are accessible in Snyder’s or Snyder Gully come from the line of concretions 3 m (9.8 ft) below the at the southern end of Canandaigua Lake, above NY Rte. Bluff Point Siltstone; the horizon lies between two 80 mm 21, 0.3 mi (0.5 km) south of Woodville (Ontario County). (3 in)-thick black shales that define the first cycle below the Bed 46c/1, with Koenenites beckeri n. sp., is a 0.22 m (8.5 cycle with the Bluff Point bed. Goniatites in Nap-8/1 in- in)-thick Fossil Log concretionary bed 5 m (16.4 ft) below clude Koenenities beckeri n. sp., K.? cf. fasciculatus, Tornoceras the base of the Middlesex Black Shale. The horizon is 0.9 m sp., and ?Lobotornoceras aff. hassoni. Nap-8/2 is a gray (3 ft) below a 0.1 m (4 in)-thick black shale and approxi- mudrock with goniatite molds, 0.61 m (2 ft) above Nap- mately 4 m (13 ft) above where continuous exposure begins. 8/1 and just below the upper 80 mm (3 in)-thick black Locality KK, Py-16, Sunset Point Gully.–In the upper shale. part of the section in Sunset Point Gully, Keuka Lake, poorly Locality JJ, Py-7, Chidsey Point.–There is an extensive preserved Koenenites occur in Bed Py-16/3, a horizon of section of the West River Shale in Chidsey Point Creek on nodules 0.91 m (3 ft) below the Bluff Point Siltstone, and the West Branch of Keuka Lake in Steuben County, 1.7 mi poorly preserved Koenenites, Acanthoclymenia, and Tornoceras (2.7 km) south of Branchport (Yates County). Bed Py-7/1 occur in the soft shales surrounding a 0.15 m (6 in)-thick is a nodular silty mudstone with crushed molds approxi- concretionary horizon (Py-16/4) beneath a 0.1 m (4 in)- mately 4.6 m (15 ft) below the base of the Bluff Point Silt- thick black shale 5.4 m (17.6 ft) above the Bluff Point Silt- stone; this is sample-horizon 8556-SD of de Witt & Colton stone. The Bluff Point Siltstone is well exposed in the section (1978: pl. 6) and Huddle (1981: B50). No goniatites were (de Witt & Colton, 1978: table 1, pl. 4). recovered from the cycle below the one with the Bluff Point Locality LL, OV-10, Mill Creek.–In the upper part of Siltstone. Bed Py-7/2 is a Fossil Log concretionary bed with Mill Creek near Lodi (Seneca County), approximately 145 small pyritic and baritic goniatites including Koenenites beck- yd (133 m) downstream of the NY Rte. 414 bridge, the go- eri n. sp. form D, Acanthoclymenia aff. neapolitana, and niatite bed with Koenenites beckeri n. sp. in the cycle below Tornoceras, 3.4 m (11 ft) below the base of the Middlesex HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 33

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

Shale exposed near the top of the section where the creek ping out in the creek floor approximately 100 yd (91 m) up- narrows near the top of a 3 m (10 ft)-high cascade. The hori- stream of the Munson Road bridge. The 8 cm (3 in)-thick zon is 1.5 m (5 ft) above a series of three thin siltstones. Bed Bluff Point Siltstone, which caps a waterfall at a ford cross- Py-7/2 might correlate westward with the Fossil Log con- ing, lies 2.4 m (8 ft) above the goniatite bed. In the section cretionary bed 18a/4b, the Williamsburgh Bed of Over et of de Witt & Colton (1978: pl. 3), the cycle that includes al. (1999, 2003), in Beards Creek (Loc. 18a) in the Genesee the Bluff Point Siltstone is in the center of the eastward- Valley. thinning wedge of West River Shale; the black shale above Locality MM, Gen 1a, Sheldrake Creek.–In Sheldrake Bed Gen 1a/1 is the lower of the two black shales shown in Creek, on the western side of Cayuga Lake, Bed Gen 1a/1 their section. is the goniatite bed with Koenenites beckeri n. sp. in the cycle Locality NN, I-3, Coy Glen.–In Coy Glen, 2.1 km (1.3 below the Bluff Point Siltstone. The horizon is a gray shale mi) west of Ithaca (Tompkins County), Bed I-3/1 is a immediately below a 0.91 m (3 ft)-thick black shale crop- mudrock with a “Naples” fauna of goniatites (Koenenites 34 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

beckeri n. sp.? and Tornoceras), bactritids, ostracods, and between the Genesee Group and succeeding strata, which styliolinids, which could correspond to the goniatite bed in in early stratigraphic classifications comprised the Portage the cycle below the Bluff Point Siltstone to the west. The and, farther east, Naples groups. The unit was formally horizon, at approximately 965 ft (294 m) elevation, is near named by Clarke (1903) for exposures near Middlesex the top of the Ithaca Shale and Sandstone (de Witt & (Yates County), and later Chadwick (1933) introduced the Colton, 1978: pl. 3). It is at the base of a 0.15 m (6 in)- name Sonyea for the combined Middlesex and Cashaqua. thick dark gray mudrock above a 0.10 m (4 in)-thick black De Witt & Colton (1959) recognized approximately 1.8 m shale near the top of a low falls, 2.7 m (9 ft) below the level (6 ft) of Middlesex Shale at Lake Erie, 4.9 m (16 ft) in the of the first driveway bridge that crosses the creek upstream Genesee Valley, 11.3 m (37 ft) at Canandaigua Lake, and a of the Elm Street Extension Bridge. Bed I-3/1 is close to the maximum of 22.9 m (75 ft) at Keuka Lake, east of which the level of sample-horizon 6791SD of de Witt & Colton unit thins rapidly toward Cayuga Lake. In the Seneca Lake (1978: pl. 3) and Huddle (1981: B47), which lies down- area and eastward, Sutton et al. (1970) recognized two stream of the Elm Street Extension Bridge. tongues of black Middlesex Shale separated by an eastward- Locality OO, I-8, Lick Brook.–In Lick Brook, 2.7 mi thickening wedge of silty facies, a lower Montour Shale, and (4.3 km) south of Ithaca, Bed I-8/1 is a silty mudrock with an upper Sawmill Creek Shale. The black shales of the Mid- goniatites (Koenenites beckeri n. sp.) in the Ithaca Shale and dlesex were not searched systematically for goniatites during Sandstone that could correspond to Bed I-3/1 in Coy Glen. our survey. The only published record is the specimen of Bed I-8/1, at approximately 980 ft (298 m) elevation, is at Sandbergeroceras syngonum Clarke, 1897, reported by Clarke the base of a 0.1 m (4 in)-thick silty mudrock overlying a (1898) from Snyder’s Gully, above Canandaigua Lake near 0.05 m (2 in)-thick siltstone and a 0.1 m (4 in)-thick black Woodville (Ontario County; Loc. 46c). H. S. Williams col- shale that forms a recess in the section on the western side lected a specimen (NYSM 12026) referable to S. syngonum of the creek approximately 100 yd (91 m) downstream of from probable equivalents of the upper Middlesex Shale in the pipeline crossing. The Sand Bank Road crossing is far- Tioga County near Harford Mills (Loc. 64), and we found ther downstream. The black shale could be the one illus- loose fragments of the species (NYSM 16577, 16578) at trated by de Witt & Colton (1978: pl. 3), 47 m (155 ft) that locality. below the base of the Middlesex Shale. Cashaqua Shale SONYEA GROUP James Hall (1840) proposed the name Cashaqua Shale for In its type area in the Genesee Valley and westward, the the distinctive green shales above the “Genesee Slate” ex- Sonyea Group (Chadwick, 1933; Rickard, 1975) comprises posed along Cashaqua Creek (now Keshequa Creek) above the black Middlesex Shale and overlying green Cashaqua Sonyea (Livingston County). This is the type locality of Shale and is perhaps the most clearly defined of the major Manticoceras sinuosum (Hall, 1843), a form that marks the cycles in the New York succession. West of Canandaigua establishment in New York of Devonian goniatites compa- Lake, the uninterrupted green shale facies of the Cashaqua rable to the assemblages of Europe. It is in the Cashaqua contrasts sharply with the interbedded dark gray and black Shale that the molluscan Naples Fauna (Clarke, 1898, 1904) shales of the Genesee Group and the microcyclothems of achieves its richest development, especially in the region of black shale, gray shale, and turbidite-siltstones in the suc- Naples (Ontario County) at the head of Canandaigua Lake. ceeding West Falls Group. East of the Genesee Valley, the The unit thickens eastward from approximately 12.8 m (42 Cashaqua Shale interfingers with and comes to overlie the ft) at Lake Erie to approximately 46 m (150 ft) in the Gene- westward-thinning wedge of turbidites comprising the Pul- see Valley and then thins eastward to 15.2 m (50 ft) near teney Shale (Colton & de Witt, 1958) and Rock Stream Canandaigua Lake. Farther southeast, between there and Siltstone (Bradley & Pepper, 1938). The Cashaqua Shale is Seneca Lake, the Cashaqua is locally condensed and ranges overlain by the Rhinestreet Shale, the basal black shale of in thickness from 1.2 to 13.7 m (4 to 45 ft) (Colton & de the West Falls Group. Sutton et al. (1970), Bowen et al. Witt, 1958; Kirchgasser, 1965) (Text fig. 10). (1974), and MacDonald (1975) have reported on the shelf Following the lithostratigraphic studies of Colton & de and near-shore facies and communities of the Sonyea Witt (1958) and Sutton (1960), Kirchgasser (1965, et seq.) Group. investigated the Cashaqua faunas, particularly the goniatites and conodonts, in sections between Lake Erie and Keuka Middlesex Shale Lake (Text-fig. 10). Several thin black shales (designated A Throughout western New York, the black Middlesex Shale through F) in the lower and upper parts of the unit and a (“lower black band” of older reports) marks the boundary few distinctive carbonate horizons provide useful markers HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 35

for locating goniatite horizons. In sections west of the Gene- black shales (D through F) (Upper Cashaqua of Kirchgasser, see Valley, dark gray shales with septarian nodule horizons 1975). Goniatites are rare in this interval but horizons with and black shales A and B overlie the black shales of the Mid- Prochorites alveolatus (Glenister, 1958) occur at the base or dlesex. This lower dark shale facies of the Cashaqua thickens just below at Murder Creek (Bed 34/19), Little Tonawanda eastwardly and in the Genesee Valley and beyond, interfin- Creek (Beds 35/17 and 35/18), Beards Creek (Bed 38/10), gers with the Pulteney/Rock Stream turbidite wedge. In east- and Hicks Point (Bed 45/17). Near the top of this dark shale ern sections, the stratigraphic positions of marker black shale facies and just below the base of the Rhinestreet Shale is the B and the Parrish Limestone above the top beds of the Rock Shurtleff Septarian Horizon (Kirchgasser, 1975), a line of Stream Siltstone indicate that all but the lowermost beds of dark styliolinid septaria well known for its rich baritized the Cashaqua were deposited after major Sonyea turbidite fauna in the area in the vinicity of Conesus and Honeoye deposition had ceased. Goniatites are rare in the Cashaqua lakes. At this level, Manticoceras sinuosum sinuosum and below marker black shale B, but an important baritized Tornoceras uniangulare obesum Clarke, 1897, are the most fauna with Manticoceras sinuosum clausium n. ssp. occurs at abundant goniatites in a fauna that includes Acanthocly- Randall Gully (Bed 44/3). This earliest Cashaqua goniatite menia neapolitana, Prochorites alveolatus, and Aulatornoceras fauna reappears, again baritized, in Whestone Brook (Bed eifliense. The upper Cashaqua marks the recurrence of the 42/5), 6.7 m (22 ft) above the marker black shale. dark shale facies with Pterochaenia seen earlier in the Penn In western sections, the lower dark shale interval of the Yan and especially West River Shales of the Genesee Group. Cashaqua is succeeded by a monotonous sequence of soft The counterpart of the Shurtleff Septarian Horizon in the olive green shales with numerous concretionary horizons. Genesee Group is the Beards Creek Horizon, a line of sep- Between Lake Erie (Loc. 30) and Murder Creek (Loc. 34), tarian nodules (Bed 18a/5) near the top of the West River Probeloceras lutheri (Clarke, 1885) is particularly abundant Shale, best seen in Beards Creek, 2.4 m (8 ft) below the base in the concretion horizons in the middle of the series. Here of the Middlesex (Text-fig. 9). also Aulatornoceras eifliense (Steininger, 1849) occurs locally. The concretionary facies of the Cashaqua extends eastward Localities.–Locality numbers with asterisks have sections il- into the thick Cashaqua section in the Genesee Valley but lustrated in Text-fig. 10. here goniatites are uncommon and less well preserved. Locality 30*, Eighteenmile Creek.–A complete section Southeast of Honeoye Lake, the concretionary facies passes of Cashaqua Shale, one typical of the area west of the Gene- into a condensed sequence of soft nodular green shales and see Valley, is exposed along Eighteenmile Creek below the silty shales (Rye Point Shale of Sutton, 1960) and in this village of North Evans (Erie County), between the railroad presumably shallower water facies the Naples Fauna attains overpass and the NY Rte. 20 bridge. The lower dark shale fa- its greatest abundance and diversity, particularly among the cies is accessible in the southern bank where exposures begin bivalves Lunulicardium and Ontaria. In this area goniatite approximately 200 yd (183 m) upstream of the railroad faunas dominated by Manticoceras sinuosum sinuosum are lo- crossing, and there a few goniatites were recovered from cally common in the nodule beds and thin current-reworked Beds 30/3 and 30/4. Marker black shale B, which caps a line limestones, especially the nodular red and green Parrish of concretions (Bed 30/5) at the top of the dark shale inter- Limestone (Clarke & Luther, 1904) near Naples. As noted val, 2.2 m (7.25 ft) above the Middlesex Shale, extends east- by Clarke (1885), the nodular facies of the Cashaqua is re- ward to the Genesee Valley and possibly as far east as Randall markably similar to the Kramenzelkalk facies of the Euro- Gully (Loc. 44) in Ontario County. The lower Cashaqua pean Upper Devonian, a condensed facies of cephalopod and contact with the underlying Middlesex Shale were illus- limestone that formed on tectonic submarine highs or trated by Over et al. (2003: fig. 1). Schwellen (Rabien, 1956; Tucker, 1973; Krebs, 1979). The soft olive-green shales of the middle Cashaqua are Kirchgasser (1965, et seq.) has argued that the nodular facies accessible in the cliffs along the northern bank of the creek of the Cashaqua accumulated over Schwellen developed on opposite North Evans. Within these shales are prominent the Rock Stream Siltstone. In New York, this distinctive horizons of argillaceous concretions (Beds 30/7-9) with fre- nodular (Knollenkalk) facies reappears in similar strati- quent but generally poorly preserved goniatites (mostly Pro- graphic settings in the West Falls Group, again where wedges beloceras). Small specimens of Manticoceras sinuosum of turbidite siltstones at the slope margin penetrate seaward clausium n. ssp.? are common in thin lenses within Bed into the gray shale facies of the basin (Text-fig. 3). 30/10, a concretion horizon at creek level at the base of the Across the outcrop belt, the upper 3.0 m (10 ft) of the section in the steep south bank downstream of the NY Rte. Cashaqua consists of a series of dark gray shales, with the 20 bridge. The carbonate horizons above Bed 30/10 are gen- bivalve Pterochaenia locally abundant, interbedded with thin erally unfossiliferous. The dark shale interval of the upper 36 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

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. HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 37 38 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

Cashaqua is here approximately 2.7 m (9 ft) thick and in- of concretionary horizons (Beds 32/6–7 and especially cludes three thin marker black shales. The lowest of these 32/8–9) in approximately the middle of the section. These (D) extends eastward to Little Tonawanda Creek (Loc. 35) beds crop out in the bed of the creek downstream of the at Linden (Genesee County), and the next highest (E) as far point where the stream bends abruptly to the west below east as Briggs Gully (Loc. 43), Honeoye Lake (Ontario Bed 32/10, a thin but prominent limestone that forms a County). At 0.9 m (3 ft) above marker bed E and 0.9 m (3 pavement in the creek floor. This same interval of goniatite- ft) below the base of the Rhinestreet Shale is the Shurtleff rich concretions is seen in all sections between Lake Erie Septarian Horizon (Kirchgasser, 1975), a horizon that can (Loc. 30) and Murder Creek (Loc. 34). A single specimen of be traced approximately 81 mi (130 km) eastward to Con- Prochorites alveolatus was recovered from Bed 32/16, a hori- klin Gully (Loc. 47), near Naples (Ontario County). zon of large concretions 0.3 m (1 ft) below marker black Locality 30a, Pike Creek.–Approximately 6.1 m (20 ft) shale D at the base of the dark shale interval of the of the upper Cashaqua Shale is exposed in the bed of Pike Cashaqua. The lower beds of the Rhinestreet Shale are ex- Creek north of Lake Shore Road, 1.3 mi (2.1 km) northeast posed in the face of the falls below the road culvert. of Jerusalem Corners (Erie County). Fragments of goniatites Locality 33*, Cayuga Creek.–A long section through the were collected at several levels, notably in Beds 30a/1 and middle and upper Cashaqua Shale is exposed in Cayuga 30a/2 (the first concretions above where exposures begin), Creek west of Cowlesville (Wyoming County), with expo- Bed 30a/5 (a thin limestone 0.9 m or 3 ft immediately sures beginning at approximately 876 ft (267 m) elevation. below marker black shale D), and the shale horizons (Beds Goniatites are especially common in a series of concre- 30a/8, 9) immediately below and above marker black shale tionary levels (Beds 33/4 to 33/7) through a 1.2 m (4 ft)- E, 1.5 m (5 ft) below the base of the Rhinestreet Shale, thick interval of shale that crops out in the floor of the which caps a low falls. stream near the northern bank, approximately 430 yd (0.4 Locality 31, Smoke Creek.–Along the South Branch of km) downstream of the point where the Erie/Wyoming Smoke Creek below California Road, 1.6 mi (2.6 km) county line crosses the creek and below where NY Rte. 354 northwest of Orchard Park (Erie County), a complete sec- turns abruptly from southeast to east. The most productive tion of Cashaqua Shale crops out 0.5 mi (0.8 km) upstream horizon is Bed 33/5 in which pyritic goniatites (especially of the NY Rte. 20 bridge. The sequence is similar to that Probeloceras lutheri) are associated with plant fragments (fos- seen at Eighteenmile Creek (Loc. 30). Goniatites were not sil logs). noted in the septaria and dark shales of the lower Cashaqua Higher and less fossiliferous units of the Cashaqua and exposed near stream level but Probeloceras lutheri is common the contact with the Rhinestreet Shale are well exposed up- in the series of prominent concretionary horizons and sur- stream along the southern bank near the county line. There rounding green shales exposed in the steep embankment Bed 33/8 is the thin concretionary limestone exposed near (Beds 31/4 through 31/7). These units correlate with Beds the water level and Bed 33/9 is the prominent limestone at 30/7 to 30/10 at Eighteenmile Creek. The lowest of these the base of the embankment. Bed 33/15 is the prominent (Bed 31/4) is 2.6 m (8.5 ft) above marker black shale B. The ledge of limestone 1.7 m (5.5 ft) below marker black shale concretions of Bed 31/5 yield P. lutheri and Aulatornoceras D. Bed 33/12 outcrops at stream level where a southward- eifliense. flowing tributary enters from the north at approximately Near the top of the embankment and in the southwest- 902 ft (274 m) elevation. erly flowing side creek, poorly preserved goniatites were re- Locality 34*, Murder Creek.–Several goniatite horizons covered from two horizons in the dark shale facies. Bed 31/2 crop out in the 24.7 m (81 ft)-thick section of Cashaqua lies immediately above marker black shale D and Bed 31/14 Shale in the ravine of Murder Creek, above Griswold, 1.5 mi is 0.3 m (1 ft) below the base of the Rhinestreet Shale. (2.4 km) south of Darien City (Genesee County). The Locality 32*, Cazenovia Creek.–Approximately 15 m Cashaqua/Rhinestreet contact forms the crest of the falls (50 ft) of Cashaqua Shale are exposed in the ravine of a west of Chick Road at approximately 1,122 ft (342 m) ele- northwesterly flowing tributary to Cazenovia Creek, be- vation. The lowermost 6.1 m (20 ft) of Cashaqua Shale is tween the main stream and the intersection of North Davis only intermittently exposed. Goniatite records begin above and Conley Roads, 0.8 mi (1.3 km) south of Spring Brook this interval, in the concretions of Bed 34/3, which underlies (Erie County). Goniatites were noted in Bed 32/4, a concre- marker black shale B. Bed 34/5 with Manticoceras sinuosum tion horizon 1.5 m (5 ft) above the 75 mm (3 in)-thick sinuosum is a horizon of enormous concretions, to 1.1 m marker black shale B, and in Bed 32/5, 1.5 m (5 ft) higher, (3.5 ft) long, 1.7 m (5.5 ft) above the black shale; Bed 34/6, but the best collections (with Manticoceras sinuosum, Probe- with Aulatornoceras eifliense is a smilar horizon with lenses of loceras lutheri, and Aulatornoceras eifliense) come from a series microfossils, which is well exposed in the creek floor, 1.7 m HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 39

(5.5 ft) above Bed 34/5. Upstream is Bed 34/7, the most creek approximately 3.7 m (12 ft) above where exposures productive horizon (with M. sinuosum sinuosm and Probelo- begin and 4.9 m (16 ft) below the top of a concrete retaining ceras lutheri) in the section, a layer of disk-shaped concre- wall bordering the southern side of the road at the edge of tions with unusual, knobby upper surfaces, which crops out the stream. A thin black shale noted by Colton & de Witt at the base of the western embankment in which Beds 34/8 (1958) approximately 0.9 m (3 ft) below the goniatite hori- through 34/10 are prominently exposed. zon was not seen. Goniatites (Probeloceras lutheri) also occur Bed 34/12 is a horizon of variably sized concretions with in the concretions of Bed 37/7 (Loc. 3008-7 of Kirchgasser, rich microfossil lenses and common goniatites (Manticoceras 1975) and in Bed 37/8 (with M. sinuosum sinuosum and P. sinuosum sinuosum and Probeloceras lutheri), which crops out lutheri), 1.7 m (5.5 ft) and 1 m (3.25 ft) below the top of the in the bed of the stream just below the point where a west- wall. erly flowing side creek joins the main stream below the right Across the road from the concrete wall is a succession of angle bend in Chick Road (1,168 ft or 356.0 m elevation goniatite horizons (Beds 37/9 through 37/15) through a 3 mark). Bed 34/13 is a 76 mm (3 in)-thick limestone-concre- m (10 ft)-thick interval in the lower part of the lower of two tion horizon at the confluence. Near the top of the green- roadcuts along East Bethany Road. This interval of relatively shale interval of the Cashaqua, goniatites are present but abundant goniatites is the easternmost expression of the uncommon in Bed 34/17, a thin limestone with Aula- mid-Cashaqua goniatite “zone” noted in sections farther tornoceras outcropping at the base of the falls. Above, Bed west. Bed 37/10 is the prominent limestone near the bottom 34/19 is a layer of large concretions with occasional goni- of the embankment and Probeloceras lutheri is especially atites (Prochorites alveolatus) 0.69 m (2.25 ft) below marker abundant in the underlying meter of shales (Bed 37/9). Beds black shale D. A few goniatites (?Probeloceras lutheri) were 37/11-12, with Manticoceras sinuosum sinuosum and P. also recovered from the dark shales of Bed 34/21 beneath lutheri, are a pair of concretionary horizons 0.6 m (2 ft) marker black shale E. above Bed 37/10; Bed 37/13 is a concretion horizon with P. Locality 35*, Little Tonawanda Creek.–A complete sec- lutheri and Tornoceras uniangulare in the surrounding shales, tion of Cashaqua Shale crops out in a tributary to Little 0.76 m (2.5 ft) higher, and Bed 37/14 is a shale horizon with Tonawanda Creek below the railroad tracks approximately P. lutheri and Aulatornoceras, 0.9 m (3 ft) above. Bed 37/15 1.3 mi (2.1 km) northwest of Linden (Genesee County). is a concretionary level, with T. uniangulare obesum, 0.61 m Goniatite records (Probeloceras lutheri) begin in approxi- (2 ft) above Bed 37/14. mately the middle of the section in Bed 35/5, a thin but Locality 38*, Beards Creek.–The upper 12.2 m (40 ft) prominent limestone-concretion horizon 2.4 m (8 ft) below Cashaqua Shale is exposed in the main channel of Beards the entry of a side creek from the southeast. More produc- Creek, 0.64 km (0.4 mi) northwest of Pine Tavern Corners tive goniatite horizons are in the concretions of Bed 35/7 at (Livingston County). A diverse, well-preserved goniatite the entry of the side creek and especially Bed 35/8, with fauna was collected from pyritic green shales (Bed 38/2) sur- Manticoceras sinuosum and P. lutheri, 0.3 m (1 ft) higher. rounding a discontinuous horizon of concretion and nod- Upstream, the prominent limestone-concretion layer at the ules in the low southern bank of the creek, 1.7 m (5.5 ft) base of the main western embankment is Bed 35/9. In the above where continous exposure begins and 10.5 m (34.5 ft) upper Cashaqua, pyritic specimens of Prochorites alveolatus below the Cashaqua/Rhinestreet contact. Bed 38/2 yields occur in the dark shales (Beds 35/17-18) just below and Manticoceras sinuosum sinuosum, Probeloceras lutheri, and above marker black shale D. Aulatornoceras. In the upper Cashaqua, Acanthoclymenia Locality 37, Wyoming Gulf.–The middle and upper neapolitana was recovered from dark gray shales (Bed 38/10) Cashaqua Shale are exposed in Wyoming Gulf along Village beneath marker black shale E, and a few goniatite fragments Brook and in roadcuts on the northeastern side of East were noted in the Shurtleff Septarian Horizon (Bed 38/11). Bethany Road, above the village of Wyoming (Wyoming Locality 39*, Buck Run Creek.–The Cashaqua Shale at- County). The Cashaqua/Rhinestreet contact is exposed in tains its greatest thickness, approximately 45.7 m (150 ft), the roadcut above the intersection with Wass Road. Goni- in the Genesee Valley. The best section for collecting is along atites are scattered through the approximately 24.4 m (80 ft) Buck Run Creek, southwest of Mount Morris (Livingston of section but two horizons produce the best faunas. Bed County), beginning several hundred yards (meters) west of 37/5 is a distinctive horizon of very large concretions (to 0.6 NY Rte. 36. In the lower part of the succession, a 6.4 mm m or 2 ft long) with abundant wood fragments, lenses of (0.25 in)-thick black shale, 5.8 m (19 ft) above the top of microfossils, pyrite, and occasionally well-preserved goni- the Middlesex Shale and 0.6 m (2 ft) below a horizon of un- atites, including Manticoceras sinuosum and Tornoceras uni- usual, small, oddly shaped concretions, is marker black shale angulare obesum. The horizon crops out in the bed of the A. Four meters (13 ft) above this black shale is a layer of 40 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

small, very dark gray, egg-shaped concretions (Bed 39/3) gulare obesum. The Cashaqua/Rhinestreet contact is at ap- and goniatites (Manticoceras sinuosum) are first seen in the proximately 1,200 ft (366 m) elevation and the septarian concretions of Bed 39/4, which lies 0.6 m (2 ft) above. bed lies 1.8 m (6 ft) below. Marker black shale C is a 2.5 cm (1 in)-thick, prominent Locality 41a, North McMillan Creek.–The upper 6.1 m horizon in approximately the middle of the section in the (20 ft) of Cashaqua Shale is exposed above East Lake Road, southern embankment in the vicinity of a normal fault. Beds in North McMillan Creek on the southeastern side of Cone- 39/11 and 39/13 are concretion horizons with M. sinuosum sus Lake (Livingston County). Baritized goniatites are com- and Aulatornoceras, 1.5 m (5 ft) and 0.61 m (2 ft), respec- mon in the Shurtleff Septarian Horizon (Bed 41a/5), which tively, below this black shale. Beginning 4.6 m (15 ft) above crops out on the northern bank just above creek level, 1.5 m the black shale and continuing for 6.1 m (20 ft) is a succes- (5 ft) below the Cashaqua/Rhinestreet contact, which is at sion of concretionary horizons (Bed 39/16 through 39/20) approximately 880 ft (268 m) elevation. with occasional goniatites, with the best collections coming Locality 42*, Whetstone Brook.–A complete section of from pyritic shales surrounding Bed 39/17 (with Probeloceras Cashaqua Shale is seen in Whetstone Brook above US Rte. lutheri and Aulatornoceras eifliense ) and Bed 39/19 (with M. 20A, 2 mi (3.2 km) west of Honeoye (Ontario County). sinuosum sinuosum). The section commences above the top of the Rock Stream The valley narrows near the top of the section and there Siltstone, which caps the first falls south (upstream) of the the dark shale facies of the upper Cashaqua is exposed in the highway. Marker black shale B is 3.7 (12 ft) above the con- face of a falls, and above that, the Cashaqua/Rhinestreet tact and 6.7 m (22 ft) higher, baritized goniatites, including contact is at the top of a narrow flume. Acanthoclymenia Manticoceras sinuosum clausium n. ssp. and Tornoceras unian- neapolitana is found in the 0.6 m (2 ft) of dark shales (Bed gulare obesum, are common in lenses within Bed 42/5, a 39/22), 0.3 m (1 ft) below marker black shale E, and above, horizon of enormous concretions in the floor of the creek. baritized specimens (including Manticoceras sinuosum sinu- Upstream, goniatites are less common in similarly large con- osum) were recovered from the Shurtleff Septarian Horizon cretions of Beds 42/6-7, 0.46 m (1.5 ft) and 1.5 m (5 ft) (Bed 39/24), 1.7 m (5.5 ft) above the black shale and 1.4 m above Bed 42/5. Bed 42/9 is a distinctive 25 mm (1 in)- (4.5 ft) below the base of the Rhinestreet Shale. thick brick-like limestone exposed just below the platform Locality 40*, Cottonwood Point Gully.–Approximately or floor of the stream at the base of the high falls. In the face 30 m (100 ft) of Cashaqua Shale is exposed in the main of the falls, a few goniatites were noted in the concretions of ravine west of NY Rte. 256 above Cottonwood Point (Liv- Bed 42/13, 8.2 m (27 ft) above Bed 42/9. Near the top of ingston County), on the western side of Conesus Lake. Bed the falls, the Shurtleff Septarian Horizon (Bed 42/18) lies 40/1 is a horizon of large concretions in the northern bank 1.4 m (4.5 ft) below the base of the Rhinestreet Shale. No at the edge of an open field, just above where exposures goniatites were recovered from the septarian nodules in begin. Here Manticoceras sinuosum, Probeloceras lutheri, and place, but a loose block with a rich fauna found near the Tornoceras uniangulare obesum are found in the concretions base of the Cashaqua section is believed to be from the hori- that envelop fossil logs. Goniatites are sporadically distrib- zon (locality 3004 of Kirchgasser, 1975). uted through succeeding horizons but near the top of the Locality 43*, Briggs Gully.–The Cashaqua Shale is ex- section, baritized material (with M. sinuosum sinuosum, posed in ravines above East Lake Road in the area of Briggs Acanthoclymenia neapolitana, and T. uniangulare obesum) is Gully, on the southeastern side of Honeoye Lake (Ontario concentrated in Bed 40/13, the Shurtleff Septarian Horizon, County). The lower Cashaqua is exposed in the third stream exposed in the face of the falls, 2.1 m (7 ft) above marker north of Briggs Gully and there a few goniatites were recov- black shale E and 1.1 m (3.75 ft) below the base of the ered from Bed 43/2, 3.0 m (10 ft) below marker black shale Rhinestreet Shale. B. In Briggs Gully, just above where outcrops begin, a rela- Locality 41, Shurtleff’s Gully.–A section through the up- tively rich fauna of baritized goniatites was found in Bed permost Cashaqua Shale crops out above Cleary Road in 43/3, a line of small concretions in the northern bank, 7.9 Shurtleff’s Gully (Shurger’s Glen), a tributary to Kinney m (26 ft) above the marker black shale and 2.4 m (8 ft) Creek, 1.5 mi (2.4 km) southeast of Livonia (Livingston above a thin but prominent siltstone. Bed 43/3 is believed County). This is the type locality of the Shurtleff Septarian to be the type locality of Manticoceras sinuosum tardum Horizon (Bed 41/1), which in this area contains the mar- Clarke, 1898, which occurs with Probeloceras lutheri, vellously preserved baritized assemblage originally described Tornoceras uniangulare obesum, and Aulatornoceras. Near the by Clarke (1898, 1899a, b). Included are Manticoceras sin- top of the section, fragments of baritized goniatites were uosum sinuosum, Prochorites alveolatus, Acthoclymenia noted in the Shurtleff Septarian Horizon (Bed 43/13), 1.8 neapolitana, Aulatornoceras eifliense, and Tornoceras unian- m (6 ft) below the Cashaqua/Rhinestreet contact. HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 41

Locality 44*, Randall Gully.–Two major goniatite levels (15 ft)-high waterfall marks the Rock Stream/Cashaqua con- are seen in the Cashaqua Shale of Randall Gully, 2.5 mi (4 tact. Goniatite fragments were noted in the nodular Parrish km) south of Bristol Center (Ontario County). The section Limestone (Bed 45/2), 9.4 m (31 ft) above the contact and commences below Dugway Road at approximately 1,120 ft 1.8 m (6 ft) above the point where the northern and south- (341 m) elevation where a 0.1 m (4 in)-thick siltstone at the ern branches join at approximately 1,080 ft (329 m) eleva- top of a low falls marks the top of the Rock Stream Siltstone. tion. Higher up, in the southern branch, 1.5 m (5 ft) above A pair of thin siltstones mark the top of a 4 m (13 ft)-thick the Parrish Limestone, Manticoceras sinuosum, Probeloceras interval of silty lower Cashaqua Shale and 2.4 m (8 ft) above lutheri, and Tornoceras occur in the shales in the vinicity of is Bed 44/3, a 38 mm (1.5 in)-thick, discontinuous concre- the concretions of Bed 45/4 and in the shales of Bed 45/6, tionary lens with clusters of baritized goniatites, mostly 1.8 m (6 ft) above. A 0.9 m (3 ft)-thick interval of nodular Manticoceras sinuosum clausium n. ssp. and Tornoceras unian- shales begins 1.8 m (6 ft) above Bed 45/6, and 1.2 m (4 ft) gulare obesum. Marker black shale B, with a rusty weathering and 2.4 m (8 ft) above the nodular interval are Beds 45/11 shale below, lies 3.2 m (10.5 ft) above Bed 44/3. Higher in and 45/13, respectively, thin limestones with goniatite frag- the section, a side stream enters from the north at approxi- ments. In the upper part of the section, goniatites, including mately 1,170 ft (356 m) elevation and 1.7 m (5.5 ft) above Prochorites alveolatus, M. sinuosum, P.lutheri, and Tornoceras, is a thin but prominent siltstone. The goniatites M. sinuosum are common in the shales surrounding the nodules of Bed sinuosum, Probeloceras lutheri, and T. uniangulare obesum are 45/17 (locality 3014-17 of Kirchgasser, 1975), 1.5 m (5 ft) particularly common in Bed 44/4, a 50-100 mm (2-4 in)- below the Shurtleff Septarian Horizon (Bed 45/18, with M. thick argillaceous limestone lens that outcrops 2.3 m (7.5 sinuosum sinuosum), which lies 2.3 m (7.5 ft) below the ft) above the siltstone. Above Bed 44/4, a few goniatites were Cashaqua/Rhinestreet contact; some of the septarian nod- noted in the concretions of Beds 44/5 and 44/6 outcropping ules at this locality contain abundant plant and fish frag- just downstream of the covered interval, which continues to ments and, in places, baritized goniatites. the base of the high falls, at the top of which is the Locality 46, Cook Point.–A 20 m (65 ft)-thick section of Cashaqua/Rhinestreet contact. Cashaqua Shale is found in the ravine above Cook Point, Locality 44a, Barnes Gully.–Two goniatite horizons were Canandaigua Lake, which crosses NY Rte. 21 at the edge of noted in the discontinuous exposure of the Cashaqua Shale Bristol Springs (Ontario County). The succession com- in Barnes Gully above Foster Point, Canandaigua Lake (On- mences above the high falls capped by a 0.13 m (5 in)-thick tario County). In the southern branch of the gully, east of siltstone that marks the Rock Stream/Cashaqua contact. The NY Rte. 21, three small tributaries that join the creek from first line of concretions above the contact is Bed 46/1 and a the southwest provide convenient geographic markers. The few goniatites are seen in the succeeding 1.5 m (5 ft) of lowest enters 1.1 m (3.5 ft) above the top of the waterfall at shale. They are more common, however, in the nodules of the Rock Stream/Cashaqua contact. The second and largest Bed 46/2, the Parrish Limestone, just below a covered inter- side creek enters 2.9 m (9.5 ft) above the base of the val 5.8 m (19 ft) above the base of the Cashaqua and in the Cashaqua, and 0.76 m (2.5 ft) higher is a thin, dark gray, shales surrounding a similar nodular horizon (Bed 46/3) 1.8 pyritic shale that is believed to be black shale B. The third m (6 ft) higher. A 0.15 m (6 in)-thick siltstone lies 1.2 m (4 side creek joins 0.9 m (3 ft) above this shale horizon, and in ft) above Bed 46/3 and a few goniatites were noted in the the narrow gully of the tributary, 5.8 m (19 ft) above the shales surrounding the concretions of Beds 46/7-8, 1.2 and confluence, is a 6.4 cm (2.5 in) argillaceous limestone (Bed 2.0 m (4 and 6.5 ft) above the siltstone. The 44a/2) with few but well-preserved goniatites, especially Cashaqua/Rhinestreet contact is at approximately 1,020 ft Manticoceras sinuosum sinuosum and Probeloceras lutheri. (311 m) elevation. Near the top of the section, at the point where exposures Locality 46a, Long Point.–Goniatites are common in the begin west of NY Rte. 21, goniatites (?Probeloceras lutheri) 38 mm (1.5 in)-thick Parrish Limestone (Bed 46a/1) and occur in the thin limestone beneath a bed of nodules (Bed capping shales that crop out at approximately 980 ft (299 44a/8), 4.3 m (14 ft) below the Cashaqua/Rhinestreet con- m) elevation in the creek above Long Point, which crosses tact. NY Rte. 21, 0.4 mi (0.6 km) southeast of Bristol Springs Locality 45, Hicks Point.–A complete sequence through (Ontario County). the Cashaqua Shale is exposed in the ravine above Hicks Locality 46b, Whiskey Point.–Molds of goniatites are Point, Canadaigua Lake, below NY Rte. 21, 2.3 mi (3.7 km) common in the nodular shales of the 76 mm (3 in)-thick northeast of Bristol Springs (Ontario County). The section Parrish Limestone (Bed 46b/1) in the ravine above Whiskey commences at approximately 1,055 ft (322 m) elevation Point on the eastern side of Canandaigua Lake. The horizon where a 15.2 cm (6 in)-thick siltstone at the top of a 4.6 m crops out at approximately 980 ft (299 m) elevation, 7.9 m 42 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

(26 ft) above the top of the Rock Stream Siltstone. the Shurtleff Septarian Horizon, 0.9 m (3 ft) below the base Locality 46c, Snyder’s Gully.–At the southern end of of the Rhinestreet Shale. Canandaigua Lake, in Snyder’s or Snyder Gully, 0.4 mi Locality 47a, Rumpus Hill.–Goniatites are also common (0.64 km) south of Woodville (Ontario County), a few go- in the Parrish Limestone (Bed 47a/1) in the tributary to the niatites were recovered from the Parrish Limestone (Bed West River on Rumpas Hill, which crosses NY Rte. 245 ap- 46c/1), a 50 mm (2 in)-thick nodular horizon exposed in proximately 5 mi (8 km) northeast of Naples (Ontario the creek bed above a covered interval 3 m (10 ft) above the County). The gully is 0.1 mi (0.16 km) northeast of the top of the high falls over the Rock Stream Siltstone. The turnoff leading to Clark Gully, which is on the opposite side horizon is approximately midway between NY Rte. 21 and of the valley. The horizon consists of 12.7 cm (5 in) of nodu- Griesa Hill Road. This is locality 3015 of Kirchgasser lar shale and lies 0.9 m (3 ft) above the top of the Rock (1975). The Middlesex Shale at this locality is the source of Stream Siltstone. NYSM 4077, the lectotype, selected herein, of Sandberge- Locality 47b, Clark Gully.–On the eastern bank of Clark roceras syngonum. Gully, on the southern side of South Hill, 5.3 mi (8.5 km) Locality 46d, Griesa Hill Road.–The Parrish Limestone northeast of Naples (Ontario County), goniatites are com- (Bed 46d/1) crops out 3.4 m (11 ft) above the top of the mon in the 0.1 m (4 in)-thick Parrish Limestone (Bed Rock Stream Siltstone in the third stream crossed by Griesa 47b/1) that crops out 1.8 m (6 ft) above the top of the Rock Hill Road north of the intersection with NY Rte. 21, 3 mi Stream Siltstone. (4.8 km) north of Naples (Ontario County). This is locality Locality 47c, Lee Cemetery.–A few goniatites (Mantico- 3009 of Kirchgasser (1975). Manticoeras sinuosum sinuosum ceras sinuosum and Probeloceras lutheri) were recovered from and Probeloceras lutheri are relatively common in the Parrish the 0.13 m (5 in)-thick Parrish Limestone (Bed 47c/1) that Limestone here and at most localities in the area near crops out 1.2 m (4 ft) above the Rock Stream Siltstone in the Naples. gully crossing NY Rte. 245, 0.1 mi (0.16 km) northeast of Locality 46e, Bristol Springs Road.–The Parrish Lime- Lee Cemetery and 5.7 mi (9.2 km) northeast of Naples (On- stone (Bed 46e/1) forms a ledge in a tributary to Naples tario County). Creek that crosses NY Rte. 21, 0.9 mi (1.4 km) northeast of Locality 47d, School No. 7, Middlesex.–Manticoceras the intersection of the highway and the Bristol Springs sinuosum sinuosum is particularly common in the Parrish Road, Naples (Ontario County). Limestone (Bed 47c/1), cropping out in a stratigraphic set- Locality 46f, Bristol Springs Road Gully.–In the lower ting similar to Loc. 47b, in the large ravine 0.1 mi (0.16 part of the narrow gully above the intersection of NY Rte. km) northeast of the site of old School #7 on West Avenue, 21 and the Bristol Springs Road, Naples (Ontario County), 1.7 mi (2.7 km) southwest of Middlesex (Yates County). the Parrish Limestone (Bed 46f/1) is 76 mm (3 in) thick Locality 47e, Lower East Hill Road.–In the gully be- and lies 2.7 m (9 ft) above the top of the Rock Stream Silt- tween NY Rte. 245 and Lower East Hill Road, 0.9 mi (1.4 stone. km) south of Middlesex (Yates County), the nodular Parrish Locality 47*, Conklin Gully.–One of the most complete Limestone (Bed 47e/1) is poorly exposed but has yielded a and accessible sections of Cashaqua Shale in the Naples area few goniatites. is the 15 m (50 ft)-thick sequence above the first falls in Locality 47f, School No. 2, Italy Valley.–In the lower Conklin Gully (formerly Parrish Gully), approximately 400 part of the creek that crosses the Italy Valley Road at the site yd (0.4 km) southeast of the intersection of Rumpas Hill of old School #2, 3.2 mi (5.1 km) north of Italy (Yates (Parish) Road and NY Rte. 245, northeast of Naples (On- County), the Parrish Limestone (Bed 47f/1) is a 0.1 m (4 tario County). This is the type locality of the Parrish Lime- in)-thick nodular shale with Manticoceras sinuosum, capped stone (Bed 47/2), the “Goniatite Concretionary Layer” of by shales with abundant auloporid corals, which crops out Clarke (1885), which is the 10.2 cm (4 in)-thick nodular opposite the school site and lies 0.9 m (3 ft) above the Rock limestone outcropping along both banks of the creek, 2.1 Stream Siltstone. m (7 ft) above the Rock Stream/Cashaqua contact (Kirch- Locality 48*, Wagener Glen.–A condensed, 4.3 m (14 gasser, 1965, et seq.). Manticoceras sinuosum sinuosum is com- ft)-thick section of Cashaqua Shale is exposed in Wagener mon and Probeloceras lutheri and Tornoceras occur. A similar Glen (West Branch of Keuka Lake) above the first falls west but unfossiliferous horizon lies 2.1 m (7 ft) above Bed 47/2 of the roadbridge, 0.5 mi (0.8 km) north of Pulteney and a few goniatites were noted in the shales beneath the (Steuben County). There the 0.28 m (11 in)-thick Parrish small concretions of Bed 47/4, 0.9 m (3 ft) higher. In the Limestone (Bed 48/1) is only 76 mm (3 in) above the 0.3 m dark shales at the top of the section, baritized goniatite frag- (1 ft) ledge of siltstone that marks the top of the Rock ments were noted in Bed 47/12, the easternmost locality of Stream Siltstone. No goniatites have been collected from the HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 43

Parrish Limestone, but Probeloceras lutheri is common in a a better unit for definition of the top of the Rhinestreet bioturbated shale (Bed 48/2) beneath a 76 mm (3 in)-thick Shale. black shale 0.3m (1 ft) above. A sequence of siltstones, silty The beds of the lower two thirds of the Rhinestreet Shale gray shale, and black shale continues upstream to the in the western half of Erie County are “composed largely of Cashaqua/Rhinestreet contact. brownish-black and petroliferous shale in which an upward Locality 49, Bobell Hill.–At Bobell Hill Quarry, 4 mi gradation from black into dark brown or very dark gray at (6.4 km) southwest of Coventry (Chenango County), J. intervals of 2 inches (5.1 cm) to medium dark gray to Harrington collected a specimen of Manticoceras that is the medium-gray shale in beds from an inch (2.54 cm) to 5 ft easternmost known occurrence of the genus in New York (1.52 m) thick” (Pepper et al., 1956). State. The locality is just south of Schaffer Road, at an alti- It is in the upper part of the Rhinestreet Shale in the tude of approximately 1,540 ft (470 m), west of Bobell Road same area that the microcycles so characteristic of the over- and Bobell Hill. The stratigraphical equivalence of the level lying Angola Shale are progressively developed. The cycles with horizons to the west is uncertain. The facies in the are initiated with black or brown shale, the upper part of quarry is that of Cypricardella and the possible equivalent is which shows evidence of burrows as it passes rather suddenly the Enfield. The level would seem to fall within the Sonyea into medium-gray shale, the upper part of which shows the Group. development of calcareous, and often septarian, concretions. It is the concretionary levels that frequently produce well- WEST FALLS GROUP preserved goniatite faunas, but mostly only in the upper part This group comprises the Rhinestreet and Angola Shale For- of the formation (Text-fig. 13). The most productive hori- mations of New York State west of Varysburg (Wyoming zon is the Relyea Creek Horizon at Relyea Creek (Loc. 60/3) County), and their more clastic easterly equivalents. Raised in Wyoming County, and correlative or nearly correlative from formation status (Rickard, 1964; Oliver et al., 1969) horizons at Cazenovia Creek (Loc. 54/1), Varysburg (Loc. to group status by Rickard (1975), the division is defined as 57/1, 2), Johnson Creek (58/1), Stony Creek (61/1), and those strata between the basal black shales of the Rhinestreet Kennedy Gulf (63/3), which have yielded Sphaeromantico- Shale and the base of the black shales of the Pipe Creek Shale ceras oxy (Clarke, 1897), S. aff. oxy, S. rhynchostomum Formation (Text-fig. 2). (Clarke, 1898), Manticoceras lamed (G. & F. Sandberger, 1850), Carinoceras vagans (Clarke, 1898), Playfordites cf. tri- Rhinestreet Shale partitus (G. & F. Sandberger, 1850) (Loc. 58/1 only), Lin- This is the thickest of the Upper Devonian black shale in- guatornoceras aff. linguum (G. & F. Sandberger, 1851), and tercalations in western New York. It was recognized by Hall Tornoceras cf. typum (G. & F. Sandberger, 1851) (Text-figs (1843) as the “upper black shale.” The unit was defined by 13, 61). Clarke (1903: 23) and by Luther (1903) under the name No substantial effort has been made in this work to es- Rhinestreet Shale or “black shale” and later “bituminous tablish a detailed correlation of the lower Rhinestreet Shale shale” (Clarke, 1904: 213) or “second black band” (Clarke, (Text-fig. 11). This part of the sequence is not rich in goni- 1898: 106). The division was named for exposures near atites, and they are known from only a few localities. Rhinestreet Road, Naples, a sequence that represents only At Buck Run Creek (Loc. 65/1), for example (Text-fig. the early part of the unit farther west. 11), a fauna with Prochorites aff. alveolatus occurs associated In the detailed study of the West Falls Group by Pepper with a fossil-log concretionary bed 12.7 m (41.7 ft) above et al. (1956), the Rhinestreet Shale was measured as 59.4 m the base of the Rhinestreet Shale. The Sturgeon Point (Loc. (195 ft) thick at Eighteenmile Creek, thinning eastwardly 50) Lake Erie area is recorded as having yielded manticocer- to 20.7 m (68 ft) thick at the Mount Morris Dam outcrop atids (Clarke, 1899c), but the exact source has not been lo- on the Genesee River, and to 4.9 m (16 ft) thick in Urbana cated. Large concretions to 2.4 m (8 ft) across with Glen in Steuben County, but it was recognized that this re- manticoceratids occur in the vinicity of the northern duction resulted from facies changes. The top of the promontory of Grandview Bay, Lake Erie, at a level that Rhinestreet Shale was taken at the bed named the “Scraggy must be approximately 10.7 m (35 ft) below the Scraggy Bed” by Luther (1903: 1022-1023), a distinctive bed of Bed, that is, quite high in the sequence. The sections illus- pyritic pustulate concretions. Pepper et al. (1956) were un- trated for the lower Rhinestreet Shale and the sedimentary able to trace this bed east of Cazenovia Creek, but we were setting show that detailed lithostratigraphic work would en- able to trace the bed of black shale that overlies it much able correlation of the thin leaves of black shale that pro- more extensively (Text-figs 13-14) and doubtless additional gressively break away from the basal black shales as one goes work will carry it yet farther, because the black shale forms eastward. 44 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

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, read Loc. 65a.

The sedimentary microcycles of the upper Rhinestreet at Fox’s Point on the southern side of Dibble Bay, Lake Erie, Shale are traceable from near Lake Erie eastward to Kennedy but it is now best exposed in Big Sister Creek (Loc. 73, Text- Gulf (Loc. 63) in Wyoming County (Text-fig. 13). As Pep- fig. 15) and Cazenovia Creek (Loc. 54, Text-fig. 13). Cor- per et al. (1956) noted, the Scraggy Bed is soon lost, but the relation of this level eastward shows that when plotted on overlying black shale is a reliable marker for the top of the the chart of Pepper et al. (1956), it lies 67 m (220 ft) above Rhinestreet Shale. the base of their Kennedy Gulf section (no. 8) at the level of Luther (1903: 1023) noted the Scraggy Bed at lake level the uppermost black shale that they indicate. Thus, the cor- HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 45

relation here established shows that most of the Gardeau Shale and the Grimes Siltstone and Hatch Shale of the re- gion east of Varysburg is the time equivalent of the Lake Erie Rhinestreet 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 near Mount Morris (Text-fig. 3). The level seems to be well above the Table Rock Sandstone of Hall (1843: 391), Luther (1903: 1006), and Chadwick (1933: 96), that is the bed that forms the top of the Lower Falls (or Lower Portage of the Genesee River (Text-fig. 12), but the exact correlation has not been established. Pepper et al. (1956) did not mark the Table Rock Sandstone on their stratigraphical column (no. 12) but they described it and it is the unit that they showed 51 ft (15.5 m) above the base on their column no. 12 be- cause they indicated that this level is 395 ft (120 m) above the top of the Rhinestreet Shale in the Genesee River. The top of the Middle Falls lies 190 ft (58 m) higher and the top of the Upper Falls seems to lie immediately underneath the Nunda Sandstone on their chart. There are a few type specimens of goniatites from the time equivalents of the Lake Erie Rhinestreet Shale in the Genesee Valley region. The source of the syntypic specimen of Sphaeromanticoceras oxy (see Clarke 1898, 1899c, 1899a, 1899b: 78, pl. 2, figs 5-6; NYSM 3746) was given as “from the soft shales, 150 ft below the Portage sandstone, at the Lower 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 the Rhinestreet Shale of Lake Erie and not the Angola Shale (Miller, 1938: 95). The other specimen originally figured by Clarke (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). The section there was illustrated by Pepper et al. (1956, section no. 18) and, accepting their correlation of the sequence from the uppermost Gardeau Shale to upper Nunda Sandstone, the specimen is most likely to have come from equivalents of the Angola Shale of Lake Erie. We remeasured this section to establish that the base and top of the main falls lie at 61 m (200 ft) and 74.7 m (245 ft), respectively, above the base of 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 really comprise two groups. Localities 50-55 and 65 comprise lev- els generally referred to the Rhinestreet Shale. But most of the other localities represent horizons that are correlated here with the Rhinestreet Shale near Lake Erie on the basis of Text-fig. 12. Section at the Lower Portage Falls on the Genesee bed-by-bed correlations (Text-fig. 12), but which, litholog- River, Livingston County. 46 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

Text-fig. 13. Sections of the upper Rhinestreet Shale equivalents between Cazenovia Creek, Erie County, and Kennedy Gulf north of Warsaw, Wyoming County, with the locality numbers used in the text. HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 47 48 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

ically, are referred to the Gardeau Shale farther east. imately 120 yd (109 m) upstream of the US Rte. 20A bridge Locality 50, Sturgeon Point.–An old locality from which west of the town (Loc. 57, Text-fig. 13). A continuation up- manticoceratids have been reported (Clarke, 1899c) is Stur- ward of the section is illustrated in Text-fig. 15 (Loc. 79/80). geon Point, on Lake Erie Shore (Erie County) at the north- At the lower locality Bed 57/1 is a horizon of small concre- ern end of Grandview Bay. Giant septarian nodules to 2.4 m tions below a 12.7 cm (5 in)-thick siltstone. Bed 57/2, just (8 ft) across, exposed near Sturgeon Point, contain Mantic- over 2.1 m (7 ft) higher, is a fossil-log concretion horizon oceras sp. This locality might lie approximately 10.7 m (35 with a log of fossil wood, 0.66 m (2.2 ft) below a 0.13 m (5 ft) below the black marker band above the Scraggy Bed at in) black shale. The goniatite fauna from this level is excep- Point Breeze to the south (see Addendum). tionally rich and includes Carinoceras vagans, Manticoceras Locality 51, Big Sister Creek.–Along Big Sister Creek, lamed, Sphaeromanticoceras rhynchostomum, and Lingua- near Angola (Erie County), the upper Rhinestreet section tornoceras aff. linguum. Localities hereabouts were known continues downward from the Angola and the relevant hori- to Williams (1887) and formed his locality 472. zons are illustrated there (Loc. 73, Text-fig. 15). There are Locality 58*, Johnson Creek.–The upper part of the two levels of concretions below the Scraggy Bed that have Rhinestreet Shale is exposed above a cabin, 0.5 mi (0.8 km) yielded goniatites: Bed 51/b at 0.46 m (1.5 ft) below, and west-northwest of Earls, in Johnson Creek (Wyoming Bed 51/a at 1.37 m (4.5 ft) below. County), 200 yd (180 m) west of a disused railroad. Bed Locality 52, Eighteenmile Creek.–The good section 58/1 (the Relyea Creek Horizon) lies 0.66 m (2.2 ft) below along Eighteenmile Creek (Erie, Chautauqua, and Catta- the base of a 0.18 m (7 in) black shale above a thin siltstone raugus counties; Grabau 1898-1899; Pepper et al., 1956: that forms the foot of the main falls. The rich fauna of Bed section 1) has not been systematically searched for goni- 58/1 includes Carinoceras vagans, Manticoceras lamed, atites. Sphaeromanticoceras rhynchostomum, Playfordites cf. triparti- Locality 53*, Hampton Brook.–The Hampton Brook tus, and Linguatornoceras aff. linguum. The upward exten- section south of Hamburg (Erie County; Loc. 74, Text-fig. sion of this section (Loc. 81) is given in Text-fig. 15. 15), illustrated for the Angola Shale equivalents, continues Locality 59a*, Tannery Brook.–A section in the lower down into the upper Rhinestreet, but faunas have not been Rhinestreet Shale continues upward from the Upper collected. Cashaqua Shale (Kirchgasser, 1967: 138, loc. 7), 1.2 mi (2.3 Locality 54*, Cazenovia Creek.–The upper part of the km) west of Attica (Wyoming County; Text-fig. 11). The Rhinestreet Shale in Cazenovia Creek (Erie County) con- base of the Rhinestreet Shale is at approximately 1,100 ft tains concretions similar to those in the overlying Angola (335 m) elevation. No goniatites were found in the Shale (Loc. 75, Text-fig. 15). Manticoceras lamed, Sphaero- Rhinestreet Shale at this locality. manticoceras rhynchostomum, and Tornoceras cf. typum were Locality 60*, Relyea Creek.–Near South Warsaw in Re- collected from the concretions of Bed 54/1, 2.4 m (8 ft) lyea Creek (Wyoming County) is a locality known to Clarke below the basal black shale marker of the Angola Shale, and (1899c) and mentioned by him in his Guide and possibly 3.5 m (11.5 ft) above a prominent bed of septarian concre- Luther (1903: 1012) under the name of Gibson’s Glen. The tions (Text-fig. 13). section given here (Text-fig. 13) commences 500 yd (460 Locality 55*, Hunter Creek.–The top of the Rhinestreet m) west of NY Rte. 19 at South Warsaw at approximately is exposed 200 yd (183 m) east-northeast of a bridge east of 1,100 ft (335 m) altitude, above which a farm track crosses Colegrove at approximately 1,033 ft (315 m) altitude in the creek at a falls over a thick black shale. Upstream a rock Hunter Creek (Erie County), and a long section below is il- pool forms the most convenient marker for the goniatite- lustrated here (Text-fig. 13). The main collection here (Loc. rich levels. Bed 60/2 is approximately 0.9 m (3 ft) above the 55) came from a loose block collected below the Scraggy pool surface, and 0.46 m (1.5 ft) above is a 12.7 cm (5 in)- Bed. This was probably derived from the Angola Shale. Oth- thick siltstone that marks the top of a small falls. Bed 60/3, erwise no goniatites were collected from the section. the Relyea Creek Horizon, is above a 0.23 m (9 in)-thick Locality 56, Sheldon Creek.–North of Strykersville, in black shale, 0.84 m (2.75 ft) higher and 5.6 m (18.4 ft) Sheldon Creek (Wyoming County), at the very base of the below the Rhinestreet/Angola Shale contact. The rich fauna section exposed above glacial debris, a level of concretions of Bed 60/3 includes Sphaeromanticoceras oxy, S. aff. oxy, (Bed 56/a) 3 ft (0.9 m) below the marker black shale of the Carinoceras vagans, Manticoceras sp., S. rhynchostomum, and Angola, at approximately the level of the Scraggy Bed, Linguatornoceras aff. linguum. Bed 60/4 contains Mantico- yielded Tornoceras sp. (Loc. 78, Text-fig. 15). ceras lamed. Locality 57*, Varysburg.–The section along Tonawanda Locality 61*, Stony Creek.–At Stony Creek, Warsaw Creek at Varysburg (Wyoming County) commences approx- (Wyoming County), a good and complete section of the HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 49

equivalents of the upper Rhinestreet is exposed (Text-fig. microcyclothem above this, and Bed 63/2 is at the top of 13). The locality was known to Luther (1903: 1012) as Fall the next cycle above which the creek floor forms a long gully. Brook. The section commences where exposures begin, Locality 65*, Buck Run Creek.–Ten miles (16 km) 1,400 yd (1.3 km) southwest of the town center. The only southeast of Mount Morris (Livingston County), Buck Run goniatites located (Sphaeromanticoceras rhynchostomum and Creek exposes the Rhinestreet and Cashaqua Shale and Tornoceras cf. typum) are from Bed 61/1, a level apparently lower units. The section given here (Text-fig. 11) is in the equivalent to the productive interval at Relyea Creek. Bed upper part of the section just downstream from where the 61/1 is best approached from the railroad crossing, 2,200 Swanson Road Bridge crosses the creek. A 40 cm (1.3 ft)- yd (2 km) southwest of Warsaw at which the upper thick black shale is the top bed marked on the section given Rhinestreet Shale and lower Angola Shale equivalents are ac- here (Text-fig. 11, Loc. 65). Bed 65/1, a fossil-log locality, cessable high in the face of the spectacular waterfall (Text- has the goniatite Prochorites aff. alveolatus. The base of the fig. 13, Loc. 61; Text-fig. 14, Loc. 83). The marker black Rhinestreet lies 12.8 m (42 ft) below Bed 65/1 at an altitude shale at the base of the Angola Shale equivalents is 10.9 m of approximately 755 ft (230 m). The higher horizons (35.7 ft) below the base of the 15 cm (6 in)-thick siltstone marked (Beds 65/2 and 65/3) were collected for conodonts on which the railroad bridge rests, and 7.3 m (24 ft) below by J. W. Huddle, but goniatites have not been found in the base of a 0.3 m (1 ft)-thick siltstone that caps the main them. falls. There is a 12.7 cm (5 in)-thick siltstone 6.8 m (22.4 ft) Locality 65a*, Parrish Gully.–At the Canandaigua Lake lower, and the concretions of Bed 61/1 are within a 0.81 m meridian (Text-fig. 3), a section in the lower Rhinestreet (2.7 ft) interval below a 15.2 cm (6 in) black shale. Shale equivalents continues upward from the Cashagua Locality 62, Warsaw.–A roadcut along US Rte. 20A, Shale in Parrish or Conklin Gully (Loc. 47) in Yates County, west of Warsaw (Wyoming County) yielded goniatites 107 northeast of Naples (Ontario County; misnumbered 69 in yd (98 m) downhill from the intersection with Buffalo Av- Text-fig. 11). No goniatites were found in the Rhinestreet enue (the old US Rte. 20A) at an altitude of approximately Shale equivalents at this locality. 1,230 ft (375 m). Locality 63*, Kennedy Gulf.–There is a fine section in Equivalents of the Sonyea and West Falls Groups South Kennedy Gulf, a tributary of Little Tonawanda Creek, 0.75 and Southeast of Ithaca mi (1.2 km) south of Dale (Wyoming County). The section There is a remarkably confined area in which the genera (Text-fig. 13) commences 14 m (6 ft) above where a side Sandbergeroceras, Wellsites, and Schindewolfoceras occur in creek enters at approximately 1,273 ft (388 m) altitude. The the area south and southeast of Ithaca (Tompkins County; upper section can be reached from a north-south road lead- Text-fig. 14). The precise horizons represented by the vari- ing to Vernal. Convenient markers in the section are few, ous localities is a matter for individual discussion. Many and none are marked on the section of Pepper et al. (1956: seem relatable to the tongues of black shale that Sutton section 8). The top of the section given here is the crest of (1963) and Sutton et al. (1962) traced in the area (Rickard, the main waterfall in the section most easily reached from 1964, 1975), that is, for Middlesex Shale equivalents the the Vernal road. This appears to be a thin sandstone unit Montour and Sawmill Creek Shales, and for the Rhinestreet that Pepper et al. (1956) marked at 267 ft (81 m) below the equivalents the successive sequence of the Moreland, Dunn black Pipe Creek Shale at the top of their section. The Hill, Roricks Glen, and Corning Shales. These tongues have marker black shale taken as the base of the Angola Shale yet to be traced westward with precision into the standard equivalent lies 5.8 m (19 ft) below and appears to be the sequence. When this is done, it could be that these goniatites highest black shale they showed. At 34 m (37 ft) below the will be found to be more widely distributed. At present, it crest of the main waterfall is a small falls over a 0.18 m (7 is only the Sandbergeroceras faunas that are known farther in)-thick siltstone, and the best fauna collected (with Man- west, as remarked earlier, in the Naples and Honeoye areas ticoceras lamed and Sphaeromanticoceras rhynchostomum) is within the Middlesex and Cashaqua Shales. There is also a from the concretions of Bed 63/3, the probable Relyea possible record in the Tygart Valley (Randolph County), Creek Horizon, which outcrops 2.6 m (8.5 ft) lower, imme- Virginia (Tilton, 1929; Miller, 1938: 180). diately above the top of the first falls encountered working The records seem to comprise a sequence in which (1) up the section. The most distinctive band below, just above Sandbergeroceras syngonum occurs in equivalents of the Mid- the base of the section given here, is a 12.7 cm (5 in)-thick dlesex Shale, (2) Sandbergeroceras? enfieldense n. sp. occurs black shale with a 50 mm (2 in)-thick siltstone in the mid- in equivalents of the upper Cashaqua Shale and lower dle, a horizon seen also in Stony Creek. Bed 63/1, with Rhinestreet Shale, (3) Wellsites williamsi (Wells, 1956) and Aulatornoceras aff. eifliense, is the concretionary level in the W. tynani House & Kirchgasser, 1993, occur in lower 50 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

1934, is the holotype (NYSM 12028). The specimen is re- ported as having come from a layer containing the sponges Ithacadictya and Protoarmstrongia. The “Endfield” here is probably the equivalent of the Cashaqua Shale of the west (Rickard, 1964) and the black shale at the top of the quarries could be the equivalent of the Rhinestreet Shale. Locality 67, Fairfield Forest.–From a locality 1.8 mi (2.9 km) southwest of Speedsville (Tioga County), 0.7 mi (1.1 km) east of Fleet Road on States Lands Road at an elevation of approximately 1,640-1,660 ft (500-506 m), a specimen of Schindewolfoceras? aff. equicostatum was collected by J. W. Wells in 1963 (his locality 389) from a small quarry near a youth camp in Fairfield State Forest. This appears to be the Text-fig. 14. Map of localities with Triainoceratidae in rocks equiv- small quarry marked on the topographic map. We located alent to the Sonyea and West Falls groups in central New York no goniatites there. The horizon was recorded as being from State and the locality numbers used in the text. Geology from the Upper Cayuta Shale (Chesney Formation) approxi- Rickard & Fisher (1970). mately 400 ft (122 m) above a Tropidoleptus level. L. V. Rickard suggested (pers. comm., ca. 1985) that the level Rhinestreet equivalents, and (4) Schindewolfoceras chemu- could be between the Moreland Shale and the Roricks Glen ngensis Vanuxem, 1842,S.? equicostatum (Hall, 1874), and Shale, that is, a mid-Rhinestreet Shale equivalent. S.? aff. equicostatum occur in the equivalents of the middle Locality 68, Bald Mountain.–The source of the holotype or upper Rhinestreet Shale. of Wellsites williamsi is from beside a track 2 mi (3.2 km) southeast of Brooktondale, 7 mi (11.3 km) southeast of Localities.–It should be noted that although these localities Ithaca, and 1.7 mi (2.7 km) northeast of White Church are all included in the West Fall Group section, some lie in (Tioga County), and 1.2 mi (1.9 km) northwest of Bald equivalents of the Sonyea Group of the western part of the Mountain at an elevation of approximately 1,770 ft (539 state. m). This is the specimen referred by Wells (1956b) to Belo- Locality 64, Harford Mills.–Harford Mills (Cortland ceras. It was recorded as coming from the “lower part of the County) lies 15 mi (24 km) east of Ithaca (Tompkins Cayuta Shale (Chemungian Stage, upper Senecan Series), County). One of H. S. Williams’s (1913) recurrent Tropi- approximately 150 ft (46 m) above the Thiemella (?) danbyi doleptus Zone localities (Hd 12G/1b) provided Sandbergero- zone (= base of Cayuta)” (Wells, 1956b: 751). The level ceras syngonum said to be from the Enfield Shale. The locality would be above the Moreland Shale, but not far above. This is southwest of Harford Mills (Tioga County) at approxi- locality seems to be from along Bald Hill School Road– mately 1,270 ft (387 m) altitude, along the stream that flows Leonard Road Junction (Loc. 68). Another locality, south eastward from West Hill and crosses NY Rte. 38, 1.2 mi (2 of the junction, is Caroline (Loc. 94), Bald Hill, town of km) south of the county line. We also found specimens of Caroline, which is the source of Sandbergeroceras? enfieldense Sandbergeroceras (NYSM 16577 and 16578) in this gully at n. sp. (NYSM 12027). a similar level. L. V. Rickard suggested (pers. comm., ca. Locality 69, Owego.–From the general area of Owego 1985) that the level is below that of the Moreland Shale and (Tioga County), there are some old records of goniatites. probably just above the Sawmill Creek Shale, that is, an The locality of the holotype of Schindewolfoceras chemu- upper Middlesex Shale equivalent. ngense (Vanuxem, 1842) (NYSM 4073) was given as Locality 66, Hungerford Quarry.–The University quar- Owego, and a subsequent label added “Cayuta Shale”; an ries lie approximately 2.9 mi (4.6 km) east-southeast of the old label gave the number 2 (?8) and “-hem Quarry, near Cornell University campus, in Ithaca (Tompkins County), Owego.” The identification of this quarry has not been de- and south of Ellis Hollow Road. There are several quarries termined but there have long been quarries on the hillside approximately along the 1,200 ft (366 m) contour. The old just north of Owego. L. V. Rickard suggested (pers. comm., university quarry, or Hungerford Quarry, is the more west- ca. 1985) that they must be at levels equivalent to the erly, and best approached from Ellis Hollow Road; the Rhinestreet Shale of the west; the contact between the Ror- newer quarries are on either side of the road over the hill, 1.5 icks Glen, Beers Hill, and Moreland Members is below the mi (2 km) south of Turkey Hill. One large specimen of top of East Beecher Hill here. A Manticoceras sp. specimen Sandbergeroceras? enfieldense n. sp., collected by K. Caster in (CU G54) in the CU collections is listed as from “locality HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 51

29” on the southeastern side of Beecher Hill. Angola Shale We found horizons with goniatites on Mountain Road The Angola Shale was originally named for the sequence of or East Beecher Hill Road, which leaves Owego just north grey shales that overlies the Rhinestreet at approximately the of the railroad crossing of NY Rte. 96. The entrance of Ever- village of Angola (Erie County; Clarke, 1903: 24; Luther, green Cemetery is along the road and at 0.33 mi (0.5 km) 1903: 1019). The thickness assigned to the unit by Pepper farther up the hill, a 2 m (6 ft) bed of siltstone has large et al. (1956) was 68.6 m (225 ft) in Hampton Brook (Loc. slump structures; higher up, a fallen block yielded Mantic- 74) and 71.6 m (235 ft) in Cazenovia Creek (Loc. 75) (Text- oceras (NYSM 16579). The sandstones and siltstones there fig. 15). are commonly full of cyrtospiriferids, productids, and bi- Pepper et al. (1956) demonstrated that the Nunda Sand- valves. stone and West Hill Flags intercalate from the east in the Locality 70, Elmira.–The holotype of Wellsites tynani upper parts of the equivalents of the Angola Shale (see also (SUI 42318) was collected by Mr. M. C. Tynan of the Uni- Baird & Jacobi, 1999). They took the 0.6 m (2 ft)-thick silt- versity of Iowa in 1978 from material derived from a roadcut stone bed at West Falls on Cazenovia Creek, approximately for US Rte. 17, which cuts through the Moreland Shale, 0.1 20.1 m (66 ft) below the top of the Angola Shale there, as mi (0.16 km) east of the crossing of NY Rte. 116 and Wa- the westernmost edge of the Nunda Sandstone (this unit tercure Road, Elmira (Chemung County), and just below crops out just downstream from the bridge at West Falls at the East Church Street Quarries (which formed locality 7 an altitude of 920 ft or 280.4 m). In this part of the section, of Woodrow and Nugent’s itinerary for the thirty-fifth An- we found an underlying group of three black shales (infor- nual Meeting of the New York Geological Association; mally herein called “The Trinity”) that enabled correlation Coates, 1963) at which the junction between the Millport eastward to Relyea Creek (Loc. 82). The correlation of the and Dunn Hill units was identified. The horizon of the lo- lowest black shale of “The Trinity” is shown in Text-fig. 15. cality would seem to be below 910 ft (277 m) in altitude, the Associated with these black shales and siltstones are a num- figure quoted by Woodrow and Nugent. The completion of ber of goniatite horizons both at the localities shown (Text- the US Rte. 17 bypass has led to the sections there being fig. 15) and others. overgrown or obscured, but there are fine sections a little to The sedimentary microcycles of the Angola Shale con- the south at an easterly exit to Church Street, Elmira, where tinue those of the Rhinestreet, and in the lower part they poorly preserved goniatite remains were noted. There is a are developed particularly well as a group of units that can problem as to the horizon because Sutton et al. (1962: 392) readily be traced well to the east (Text-fig. 15). Thin silt- stated that the type section of the Roricks Glen is at a scout stones progressively enter eastward and up the succession. camp at the southern end of Elmira, and that is an upper One such thin turbidite crops out at the mouth of Farnham Rhinestreet unit. It remains to be investigated whether the Creek at Point Breeze (Loc. 72) Lake Erie Shore (above Bed stratal dip would carry the supposed Moreland Shale below 72/6, Text-fig. 15) and can be traced for approximately 45 the type section of the Roricks Glen. mi (72 km) to Johnson Creek (Loc. 81). The concretionary Locality 94, Bald Hill, Caroline.–A specimen of Sand- units in the lower part of the Angola Shale are particularly bergeroceras? enfieldense n. sp. collected by S. C. Hollister is rich in well-preserved goniatites. in the CU collections (1071K; plastotype NYSM 12027). It The Angola Shale is the source of some of the primary came from Bald Hill, town of Caroline (Tompkins County), collections. The type material of Sphaeromanticoceras rhyn- approximately 0.5 mi (0.8 km) south of Loc. 68 at an alti- chostomum (Clarke, 1898: 65, pl. 4, figs. 6-13, pl. 5) in- tude of between 1,700 and 1,800 ft (518 and 549 m). This cludes material from Big Sister Creek (Loc. 73) presumably level is not far above the Moreland Shale. from the localities later indicated by Clarke (1899c: 107) as Locality 95, North Chemung.–A single poor specimen near the crossing of the Lake Shore railroad and the creek; of Schindewolfoceras? sp. with Tropidoleptus in the CU collec- other material from Clarke (NYSM 3794-6) is labeled only tions (Bartwick collection, Locality 48 or 29?) is recorded as Angola. The type material of Carinoceras sororium from the Chemung Group of authors between Chemung (Clarke, 1898: 75, pl. 4, figs. 1-5, text-fig. 55) was said to and North Chemung (Chemung County). No further infor- come from “the vicinity of Angola, on the Lake Erie shore mation is available on this locality. and along Big Sister and Farnham creeks, Erie County.” This Locality 96, Van Etten.–A single triainoceratid collected again would indicate the concretionary levels low in the An- in 1882 by H. S. Williams (his locality LXII), CU 3308, is gola Shale but, as will be seen (Text-fig. 15), almost all the labeled as from Van Etten (Chemung County; Waverly concretionary levels of the microcyclothemic units here yield Quadrangle). No further information is available on this lo- goniatites so that the exact level is likely to remain uncertain. cality. The most productive horizon is the Point Breeze Goniatite 52 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

Text-fig. 15. Sections of the lower Angola Shale equivalents between Lake Erie and Warsaw, Wyoming County, with the locality numbers used in the text. Note: For Loc. 75, read 1 mi NE of Griffins Mills; for Loc. 80, read Loc. 79. HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 53 54 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

Bed, which has yielded Sphaeromanticoceras oxy, S. rhyncho- (Clarke, 1899c), he gave more precise localities that indicate stomum, Carinoceras vagans, Manticoceras lamed, M. aff. that the sections from which he collected are those along lamed, Playfordites cf. tripartitus, Linguatornoceras aff. lin- Big Sister Creek on the northeastern side of Angola (Erie guum, Aulatornoceras paucistriatum (d’Archiac & de County). Many goniatite-bearing horizons are present in the Verneuil, 1842), and Crassotornoceras aff. crassum (Matern, microcyclothemic succession (Text-fig. 15) and it is impos- 1931a) (Text-figs. 15, 61). sible to say from exactly which the type material came. The Scraggy Bed is exposed along Big Sister Creek ap- Localities.–Locality numbers with asterisks have sections il- proximately 90 yd (82 m) downstream from Mill Road lustrated in Text-fig. 15. bridge, Angola. The 0.46 m (1.5 ft)-thick marker black The lower part of the Angola Shale, especially in the shale, here taken as the base of the Angola Shale, crops out west toward Lake Erie, is one of the most goniatite-rich units approximately 20 yd (18 m) toward the bridge. Goniatites in New York. As has already been indicated (House, 1968; occur in the concretions of the first microcyclothem. Bed Kirchgasser, 1973), microcyclothemic units can be corre- 73/2 is in the second microcyclothem and this is the first lated quite widely (Text-fig. 15). Accordingly, the horizon line of large concretions below the Mill Road bridge. Beds suffixes given to the locality numbers for this formation are 73/2-4 are exposed upstream from the old mill dam to the given serially upward from the base (1-9), using the standard railroad crossing and are on the western side. The 60 cm (2 sequence established in Hampton Brook (Loc. 74) to the ft)-thick black shale is exposed in the cliff above. This seems level of the 0.6 m (2 ft) black shale, 18.6 m (61 ft) above the to be one of Clarke’s localities because he referred to sections black shale immediately above the Scraggy Bed. For higher below the railroad culvert. units, and where convincing correlation is not established, The higher succession is exposed upstream of the rail- the usual procedure is adopted. road bridge, especially on the eastern side. The 60 cm (2 ft)- Locality 71*, Farnham Creek (Lake Erie Shore).–Ap- thick black shale is at river level, 183 m (200 yd) upstream proximately 500 yd (460 m) south of the mouth of Farnham from the railroad bridge. In the embankment above there is (or Muddy) Creek (Erie County), the Angola Shale reap- a 76 cm (2.5 ft)-thick black shale at the base of the eighth pears after a concealed interval. Bed 71/9a is a horizon with microcyclothem at a level that projects to the level of the pyritized goniatites (including Manticoceras lamed and Aula- top of the lower arch of the bridge; 1.5 m (5 ft) below the tornoceras aff. eifliense) in a 0.6 m (2 ft)-thick band of black black shale is the same turbiditic siltstone as that exposed at and gray shales at the foot of the exposure. Bed 71/9b is a the mouth of Farnham Creek (Loc. 72). The Point Breeze line of flat concretions 0.38 m (1.25 ft) higher. Goniatite Bed (Bed 73/6) with Sphaeromanticoceras oxy, S. Locality 72*, Point Breeze (Lake Erie Shore).–On the rhynchostomum, and Carinoceras vagans, is in the microcy- shore of Lake Erie, at the southern end of Grandview Bay, clothem (number six) immediately below the siltstone bed. 3.5 mi (5.6 km) south-southwest of Angola at Point Breeze A horizon of pyritic goniatites (Bed 73/9) with Manticoceras (Erie County), beds low in the Angola Shale crop out near lamed was located 1 ft (0.3 m) above a 20 cm (8 in)-thick the point. Lake level at the northern side of the point falls siltstone unit. within the lower part of the second microcyclothem above Locality 74*, Hampton Brook.–An excellent section is the black shale succeeding the Scraggy Bed, but usually these exposed along Hampton Brook, south of Hamburg (Erie lower beds are not seen. The first horizon of large concre- County; Colton, 1956; Pepper et al., 1956: section 2). The tions comprises Bed 72/2, and the concretionary level in the black shale marker above the Scraggy Bed is at approxi- third microcyclothem comprises Bed 72/3, and this is ap- mately 760 ft (232 m) altitude, downstream of the Ham- proximately at water level on the southern side of Point burg-East Eden Road bridge, 1.1 mi (1.8 km) southeast of Breeze. The fauna of the Point Breeze Goniatite Bed (House, the confluence of Hampton Brook and Eighteenmile Creek. 1968: 1066), with Manticoceras lamed and Sphaeromantico- The first falls below the bridge are over a siltstone above a 60 ceras rhynchostomum, was collected from 120 m (130 yd) cm (2 ft)-thick black shale succeeding the ninth microcy- south of Point Breeze (Loc. 72/6) from the top of a small clothem. Prominent markers to lower microcyclothems are gully showing an anticlinal fold in the sixth microcyclothem. in the overhanging turbiditic siltstone with good bottom To the south, higher beds are concealed, but the thin tur- structures in the seventh microcyclothem and a higher 0.15 bidite in the seventh microcyclothem crops out to the south m (6 in)-thick siltstone in the eighth microcyclothem. The at the mouth of Farnham (or Muddy) Creek. richest faunas come from the Point Breeze Goniatite Bed Locality 73*, Big Sister Creek, Angola.–The classic area (Bed 74/6b), and especially a rather lower concretionary for the collection of Angola Shale goniatites by J. M. Clarke level in the sixth microcyclothem (Bed 74/6a), which con- was given solely as Angola (Erie County), but in his Guide tains Sphaeromanticoceras oxy, Carinoceras vagans, S. rhyn- HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 55

chostomum, Aulatornoceras paucistriatum, and Crassotorno- section upward to Johnson’s Falls below the Strykersville to ceras aff. crassum. Downstream a footpath crosses the creek Dutch Hollow Road. on a 15 cm (6 in)-thick black shale and the concretions of Locality 79*, Varysburg.–The section given here is a Bed 74/3 in the underlying third microcyclothem yield C. composite of exposures along tributaries of Tonawanda vagans, A. auris (Quenstedt, 1846) group, and A. paucistria- Creek near the village of Varysburg (Wyoming County) tum. (Text-fig. 15; Locality 80 read 79). The lowest beds (Beds Above the bridge the section continues up to the North 79/1-4) are exposed in the floor and at the edge of Boston-Kehe Road bridge and beyond. “The Trinity” black Tonawanda Creek, downstream of the US Rte. 20A crossing shales are exposed where a side creek joins Hampton Brook on the western side of the village. Bed 79/4 is characterized at approximately 840 ft (256 m) altitude. by occurrences of Manticoceras lamed. The section continues Locality 75*, Cazenovia Creek.–In the West Branch of upward to levels near the Point Breeze Goniatite Bed (Bed Cazenovia Creek, where the Jewettville road crosses at 79/6) in the gully of the tributary west of the village below Griffins Mills (Erie County), a siltstone in the eighth mi- a disused railroad track; Clarke (1899c: 110) reported Man- crocyclothem forms a falls beneath the bridge. There is one tioceras (= Sphaeromanticoceras) oxy from approximately the prominent thin siltstone 2.5 m (8.25 ft) below which is the level of the tracks. The upper part of the illustrated section turbidite of Farnham Creek mouth (Loc. 72). The fauna of (Beds 79/11-13) is taken from exposures upstream of the the Point Breeze Goniaite Bed (Bed 75/6) comes from a railroad track in which “The Trinity” black shales can be concretionary level below the 30 cm (1 ft)-thick black shale readily located beneath a 0.66 m (2.2 ft)-thick siltstone at beneath this unit; the fauna includes Sphaeromanticoceras the top of a falls. Massive siltstones typical of the Nunda oxy, Carinoceras vagans, S. rhynchostomum, Linguatornoceras Sandstone enter 7 m (23 ft) above the top of the falls, and aff. linguum, Aulatornoceras paucistriatum, Crassotornoceras the Pipe Creek Shale (and top of Angola) projects to a level aff. crassum, and Tornoceras cf. typum. Bed 75/4 with C. va- approximately 41.5 m (136 ft) higher. On the section of gans and Manticoceras lamed is the upper level of large flat Pepper et al. (1956: section 5), only the upper parts of this concretions lower down in the fourth microcyclothem. section were referred to the Nunda, and the lowest siltstones Locality 77*, Glade Creek.–Upstream of a bridge on the were marked as West Hill Member. It should be noted that Dutch Hollow Road, on Glade Creek, 0.7 mi (1.1 km) the marker black shale that defines the base of the Angola north of Strykersville (Wyoming County), there are falls Shale (and top of Rhinestreet Shale) is probably one of the over a sandstone, 0.8 m (2.7 ft) in thickness, at an altitude two lowest black shales marked by Pepper et al. (1956) in of approximately 1,075 ft (327 m), which has been taken as their section 5. the base of the Nunda Sandstone. This is marked at the top The middle part of the illustrated section, from the Point of the section illustrated here (Text-fig. 15), which was meas- Breeze Goniatite Bed (Bed 79/6) upward to Beds 79/9 and ured downstream successively below the new and old 79/10, a Knollenkalk horizon with Carinoceras aff. vagans, bridges. A 30 cm (1 ft)-thick black shale marks the base of Manticoceras lamed, and “The Trinity” black shales, is based “The Trinity” black shales, and Bed 77/3 is a course of con- on exposures along Stony Brook east of Varysburg, the eas- cretions 0.48 m (1.6 ft) above it. There are two levels of iest access to which is a farm track that crosses the middle of Knollenkalk siltstone below, and these comprise Beds 77/2 the section. Part of the upper section was measured in a side and 77/1, the latter with Carinoceras aff. vagans. This section branch that joins Stony Brook at approximately 1,225 ft is wholly above NY Rte. 78. (374 m) altitude. Locality 78*, Sheldon Creek.–Sheldon Creek, a tributary Locality 80, Varysburg (Gassman Road).–Within the to Buffalo Creek, crosses NY Rte. 78, 1.8 mi (2.9 km) Nunda interval above the section measured for Text-fig. 15 north-northwest of Strykersville (Wyoming County). The is a Knollenkalk horizon that has yielded a single Sphaero- top beds of the Rhinestreet Shale are exposed above glacial manticoceras rhynchostomum?. The horizon (Bed 80/14), at debris above NY Rte. 78 and below gravel quarries. The approximately 1,375 ft (419 m) altitude, is 1.8 m (6 ft) marker black shale, here taken as the base of the Angola above the old bridge foundation where the Gassman Road Shale, is exposed at approximately 995 ft (303 m) altitude. crossed Stony Brook east of Varysburg (Wyoming County). The best faunas are from two levels in the fourth microcy- The horizon, within highly burrow shales with Zoophycos clothem (Beds 78/4a and 78/4b) below a 61 cm (2 ft)-thick traces, might be the middle or upper of the three nodule black shale. The fauna of Bed 78/4b includes Manticoceras bands in the Nunda Sandstone interval shown in section 5 lamed, Aulatornoceras auris group, A. aff. eifliense, and A. of Pepper et al. (1956). paucistriatum. The Point Breeze Goniatite Bed (Bed 78/6) Locality 81*, Johnson Creek.–Approximately 2.4 mi (4 was located higher in the section. The creek continues the km) north of Varysburg, on Johnson Creek (Wyoming 56 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

Text-fig. 16. Sections of the Hanover Shale equivalents between Walnut Creek at Silver Creek (Chautauqua County) and Beaver Meadow Creek at Java Village (Wyoming County), with the locality numbers used in the text. Note: The 20 cm concretionary bed below the F/F boundary 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.

County), a long section of Angola equivalents continues concretions near the top of the sixth microcyclothem, ap- above Loc. 57. The marker black shale of the basal Angola proximately 3.6 m (12 ft) above the top of the falls over silt- is in the middle of a cascade approximately 350 yd (320 m) stones in the fourth microcyclothem. Bed 81/10 can be west of disused railroad west-northwest of Earls. The Point recognized by a 0.25 m (10 in)-thick mudrock unit with Breeze Goniatite Bed (Bed 81/6), with Sphaeromanticoceras trace fossils that forms a ledge across the creek, 120 yd (110 rhynchostomum and Crassotornoceras aff. crassum, comprises m) below the north-south Maxon (Beck) Road on the hill HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 57

above. (1956: sections 11-12), this level was not indicated, but our Locality 82*, Relyea Creek.–The section in Relyea Creek survey indicates that it is the sandstone they marked approx- (Text-fig. 15) continues the Rhinestreet sequence of Loc. 60 imately 73 m (24 ft) below the top of their section 11, and (Text-fig. 13) and commences 0.6 m (2 ft) above the base of approximately 15 m (50 ft) above the bottom of their sec- a main waterfall, 650 yd (595 m) west of NY Rte. 19, at tion 12. It lies at an altitude of approximately 910 ft (277 South Warsaw (Wyoming County). Bed 82/6, the Point m). Breeze Goniatite Bed, is approximately 7.6 m (25 ft) above The level from which the syntype of Sphaeromanticoceras the base of the marker black shale at the base of the Angola oxy (NYSM 3746) came is only approximately determined, and approximately 8.5 m (28 ft) below the top of the main as has been indicated above, but would appear to be from waterfall; the fauna includes Sphaeromanticoceras rhynchosto- the area of the Lower Portage Falls at the section indicated. mum, Manticoceras aff. lamed, Playfordites cf. tripartitus, and This probably lies within late Rhinestreet Shale equivalents. Crassotornoceras aff. crassum. Locality 87, Stony Brook Glen.–The long section in Locality 83*, Stony Creek.–There is a fine section in Stony Brook Glen, through the state park in Steuben Stony Creek (Text-fig. 15) and the section given here con- County, south of Dansville (Livingston County), was fig- tinues upward from Loc. 61 (Text-fig. 13). The basal marker ured by Pepper et al. (1956: section 18). This is a locality of black shale of the Angola is seen in the face of the main wa- Clarke (1898: 78) for Sphaeromanticoceras oxy, but the pre- terfall below the railroad bridge 2,200 yd (2 km) southwest cise level was not recorded by him. There is a level of large of Warsaw (Wyoming County). The 15 cm (6 in)-thick silt- concretions in siltstones with thin fossiliferous lenticles just stone on which the railroad bridge rests is a convenient below and downstream of the old railroad bridge, but goni- marker for Bed 83/6a, which has yielded a very fine fauna atites were not noted there. with Manticoceras aff. lamed and Sphaeromanticoceras rhyn- Locality 87a. I-390–Dansville.–The Dansville area chostomum. Another bed, 83/6b, is 0.7 m (2.3 ft) above this (Text-fig. 3) is also the source of a specimen of Sphaeroman- siltstone. Bed 83/8 is a silty Knollenkalk horizon with ticoceras oxy (NYSM 16570) from the upper West Falls Sphaeromanticoceras oxy immediately above a 18 cm (7 in)- Group (Nunda or Wiscoy Sandstone equivalents) collected thick siltstone, 3.6 m (12 ft) above the bridge foundation from loose blocks by B. Oldfield (Broome Community Col- unit. lege). The horizon is a shell-rich channel fill on the southern Locality 84*, Kennedy Gulf.–Loc. 84 is a continuation side of US Rte. I-390 (between markers 15 and 14) near into the lower Angola Shale of the section in the Upper Exit 4 (Wayland), in Steuben County, south of Dansville Rhinestreet at Kennedy Gulf (Loc. 63; Text-fig. 13), south (Livingston County). Stop A-5-4, New York State Geolog- of Dale (Wyoming County). A concretion in Bed 63/4 ical Association (Kirchgasser, Over, & Woodrow, 1994: 358) yielded a fine specimen of Spaeromanticoceras oxy collected and Stop B-9, Subcommission on Devonian Stratigraphy by J. Kralick (NYSM 16536). Bed 63/4 is 2.23 m (7.3 ft) (SDS) (Kirchgasser, Over, & Woodrow, 1997: 232). above the base of the Angola Shale (Text-fig. 13). Locality 85, Wolf Creek.–There are falls along Wolf Pipe Creek Shale and Hanover Shale Creek over the Nunda Sandstone just below the village of The black Pipe Creek Shale and gray Hanover Shale were Castile (Wyoming County). A locality approximately 250 combined as the Java Formation by de Witt (1960) but we yd (230 m) downstream of the falls yielded a goniatite fauna are following older usage and including these units as part of from grey silty mudrock that includes Manticoceras lamed.A the upper West Falls Group. Details of the lithostratigraphic number of the large specimens (?Sphaeromanticoceras) have variation in these units were worked out by Pepper & de auloporid encrustations similar to those seen on goniatites at Witt (1950) and considerable detail was given on their Varysburg in levels just above “The Trinity” black shales chart. Correlations within the Pipe Creek, Hanover, and (Bed 79/12). We were not able to demonstrate that it is the lowermost black Dunkirk shales of the succeeding Canad- same horizon, but estimates based on the regional dip sug- away Group in Erie and Wyoming counties have been re- gest it is a possibility. fined by Over (1997a, b) and Over et al. (1997). A detailed Locality 86*, Genesee Gorge.–The horizons at which go- transect of uppermost Hanover and lowermost Dunkirk cor- niatites have been located in this survey in the Genesee relations between Lake Erie and the town of Java in Gorge (Wyoming and Livingston counties) are illustrated Wyoming County was also provided by Baird & Lash in Text-fig. 12. In this section, the well-known Table Rock (1990: fig. 5). Sandstone at the Lower Portage Falls is visible. The unit was The Pipe Creek Shale forms a clearly marked basal unit named by Chadwick (1933: 96) but it was first recognized of black shale that is only 0.6 m (2 ft) thick on Walnut by Hall (1843: 391). In the column given by Pepper et al. Creek (Locs 89, 89a; Text-fig. 16), south of the village of 58 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

Silver Creek, near Lake Erie in Chautauqua County, but the conodonts, particularly in the sections in Irish Gulf (Loc. unit thickens eastward and Sutton (1963) recognized the 90) and Beaver Meadow Creek (Loc. 92) and the interpre- horizon as far east as the Waverly and Owego Quadrangles tations shown on Text-fig. 16 follow his work (Over, 1997a, (Text-fig. 1). No goniatites have been reported from the Pipe b; Over et al., 1997). Creek Shale and none were noted in this study. The Hanover Shale, a medium-gray shale with black Localities.–Locality numbers with asterisks have sections il- shale bands and siltstones especially in the upper part, is 29 lusrated in Text-fig. 16. Horizons bearing goniatites in the m (94 ft) thick on Walnut Creek (Loc. 89a), a locality that Hanover Shale are restricted to the western outcrops close to has produced the best goniatites faunas in the lower part of Lake Erie and only a few sections were measured by us the unit, mostly small Dephiceras cataphractum (Clarke, (Text-fig. 16). 1898) (Beds 89a/2-4) but including Crickites lindneri Locality 88, Eden Valley.–The South Branch of Eigh- (Glenister, 1958) from a level located by G. Kloc in the teenmile Creek passes through Eden Valley (Erie County); vicinity of Beds 89a/4 or 89a/5 (Text-fig. 16). Other loose 2.5 mi (4 km) upstream of Eden Valley where the Church specimens of C. lindneri collected by G. Klapper (NYSM Road bridge crosses the creek. Approximately 200 yd (183 16537 and 16538) and R. T. Becker (HM MBC 1943) m) upstream is the old bridge where the former road crossed probably came from the same level. Bed 89a/5 or just above beside a sawmill at approximately 800 ft (244 m) altitude. is the probable source of the specimen collected by G. Kloc The Pipe Creek Black Shale lies below. The Pipe Creek is that we earlier described as Sphaeromanticoceras aff. rickardi here approximately 1.2 m (4 ft) thick, and the overlying (House & Kirchgasser, 1993: text-fig. 56G-I). G. Kloc has basal Hanover Shale has small concretionary nodules with a collection of pyritic goniatites from the Pipe Creek- manticoceratids. Immediately above follow thick black-shale Hanover contact beneath the NY Rte. 20 bridge in Walnut units. Creek (Loc. 89a), but the collection has yet to be described; Locality 89a*, Walnut Creek.–On Walnut Creek south this pyritic level was apparently covered by creek sediment of the village of Silver Creek (Chautauqua County), there is when we did our survey. The same levels near the Pipe a long and fine section of the Hanover Shale. The black Pipe Creek-Hanover contact outcropping in Silver Creek, up- Creek Shale crops out below the NY Rte. 20 bridge and the stream of the NY Rte. 20/5 bridge in the village of Silver section figured is upstream of the bridge where the basal Creek, have yielded goniatites to collectors but this locality beds of the Hanover Shale are well exposed for several hun- was not included in our survey. The lowermost Hanover is dred yards (meters) on the eastern side of the creek. Bed the source of the cotypes of Delphiceras cataphractum and 89a/4, the most productive horizon, is at stream level 40 yd probably also of the lectotype (NYSM 4092) and syntypes (37 m) above the bridge and has a thick black shale, 0.3 m of Aulatornoceras rhysum (Clarke, 1898), which came from (1 ft) above it. Beds 89a/1-3 are best exposed approximately Beaver Meadow Creek (Loc. 92), at Java in Wyoming 400 yd (365 m) farther upstream. Beds 89a/2-4 contain Del- County. phiceras cataphractum and Bed 89a/5 yielded to G. Kloc a At a higher level in the Hanover Shale at Glade Creek fine specimen of Sphaeromanticoceras aff. rickardi and one (Loc. 91) near Strykersville in Wyoming County, L. V. of Crickites lindneri. Bed 89a/5 is probably the source of the Rickard found a specimen of Sphaeromanticoceras rickardi two loose specimens of C. lindneri described herein. (NYSM 12032) from a concretionary bed (Bed 91/7) within Locality 89*, Walnut Creek.–Higher upstream on Wal- a series of silty beds. From near the top of the Hanover Shale nut Creek in Chautauqua County, above the concealed in- in Irish Gulf (Loc. 90), ?Archoceras sp. (NYSM 16587 and terval marked by Pepper & de Witt (1950: section 1), the 16588) and ?Crickites sp. juv. (NYSM 16589) occur in Bed upper beds of the Hanover Shale are exposed. No goniatites 90/2, a level that also yields conodonts of MN Zone 13 were collected from these levels and the placing of the Frasn- (Over, 1997a, b). These are the highest known goniatites in ian/Famennian boundary is based on that by Over (1997b). the Frasnian succession in New York State. Locality 90*, Irish Gulf.–A fine section of Hanover Shale Eastward the Hanover Shale passes progressively into the commences in Irish Gulf upstream of the railroad crossing Wiscoy Sandstone (Pepper & de Witt, 1950) and at Wiscoy above where NY Rte. 391 (Boston State Road and old NY Creek (Loc. 93) goniatites have been reported but none were Rte. 219) crosses the creek, 0.3 mi (0.5 km) south of the found in this study. Luther (1911: 23) stated that Mantico- Chestnut Ridge Road and 1.6 mi (2.6 km) southeast of ceras (= Sphaeromanticoceras) oxy occasionally appears in the North Boston (Erie County; Text-fig. 16). Goniatites with Wiscoy beds. manticoceratid sutures occur in the concretions (with white The precise relations of the Frasnian/Famennian bound- barite) of Bed 90/1 (WTK 4100) in the creek floor at the ary have been elucidated by J. Over at several localities using level of the gas seeps, approximately 2.5 m above the base of HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 59

the Hanover; conveivably these could be large specimens of Bed 92/2 is probably the type horizon of Delphiceras cat- Delphiceras cataphractum but the inner whorls are not pre- aphractum and Aulatornoceras rhysum. A bed of larger con- served. The highest known goniatites in the New York cretions, 91/3, crosses the creek upstream, below the remains Frasnian occur in Bed 90/2, a 2.0-2.5 cm-thick concre- of a wooden dam. There is a considerable section exposed on tionary horizon capping a small falls in the upper Hanover the northern bank of the creek. Bed 92/4 is a horizon of Shale, 4.7 m below the base of the conodont-defined Frasn- large concretions, 0.9 m (3 ft) above stream level at the foot ian/Famennian boundary (Over 1997a, b). They consist of of this section. small, poorly preserved whorl sections of ?Archoceras sp. Locality 93, Wiscoy Creek.–Equivalents of the Hanover (NYSM 16587, 16588) and ?Crickites sp. juv. (NYSM Shale tongue into the Wiscoy Sandstone along Wiscoy 16589). This baritic crinoid and gastropod-rich horizon was Creek, Wiscoy (Allegany County), below the dam. A visit by illustrated by a concretion symbol by Over (1997b: fig. 8). the Friends of the Devonian in 1962 (J. W. Wells, pers. Locality 91*, Glade Creek.–North of Strykersville, in comm.) found goniatites near the second cascade, but none Glade Creek (Wyoming County), the middle and upper were located by us. parts of the Hanover Shale are well exposed between the Stykersville Road north to Dutch Corner and the Plants CANADAWAY GROUP Corner Road to Sheldon. The basal black Pipe Creek Shale The Canadaway Group (Chadwick, 1933) comprises the crosses the creek at approximately 1,220 ft (372 m) altitude, next sequence initiated by the entry of another prominent and the basal black Dunkirk Shale at approximately 1,350 black shale, the Dunkirk Shale, which is succeeded by the ft (411 m). In the lower part of the section a boundary fence South Wales Shale, Gowanda Shale, Laona Sandstone, and crosses the creek and a short distance above is a 0.84 m (2.75 a series of other units that have not yielded goniatites and do ft)-thick black shale marker band with a thin siltstone 2 m not concern us here. Pepper & deWitt (1951) provided de- (6.5 ft) higher. A loose specimen of Sphaeromanticoceras tailed sequences through the group and Baird & Lash rickardi (NYSM 12033) was found in this part of the lower (1990), Over (1997a, b), and Over et al. (1997) added sig- Hanover Shale. Bed 91/1 is a horizon of small nodules be- nificant refinements, particularly in the lower Dunkirk tween two thin black shales, 0.69 m (2.25 ft) apart, which Shale. Goniatites have only been discovered in the Gowanda lie above the siltstone. At 0.6 m (2 ft) above the higher thin Shale during this work at two levels, a level with Truyolsoceras black shale is a broad spread of small concretions forming a cf. bicostatum (Hall, 1843) (Loc. 99) on Walnut Creek and ledge across the stream, which constitute Bed 91/2. Bed a rather higher level that has been termed the Corell’s Point 91/4 is at the foot of the falls above. Bed 91/7, a 10-15 cm Goniatite Bed (Loc. 97) with Cheiloceras (Cheiloceras) ambly- (4-6 in)-thick Knollenkalk siltstone that produced a single lobum (G. & F. Sandberger, 1851), Phoenixites concentricus specimen of S. rickardi (NYSM 12032), forms the crest of (House, 1965), and Truyolsoceras bicostatum. The level near a 0.6 m (2 ft)-high waterfall, 1.6 m (5.25 ft) above a large Forestville which yielded T. clarkei (Miller, 1938) has not 4.5 m (15 ft)-high waterfall. Bed 91/7 is approximately 7.6 been located. When the entry of Cheiloceras in Europe was m (25 ft) below the base of the Dunkirk Shale. taken as a marker to the lowest Famennian, the recognition A track down from the road between Plants Corner and of this level in New York (House, 1962) provided the first Sheldon gives the easiest access to the upper part of the sec- convincing evidence for the location of the Frasnian/Fa- tion. The track crosses the creek almost at the Hanover/ mennian boundary, but a Global Stratigraphic Section and Dunkirk contact with a 15.2 cm (6 in)-thick hard limestone Point has now been established by the International Union band 0.6 m (2 ft) below. The Huddle Rider Bed is a thin of Geological Sciences (IUGS) for the boundary in the black shale with possible cephalopod anaptychi, 0.76 m (2.5 Montagne Noire, France (Klapper et al., 1993; House, ft) below the limestone band. Becker et al., 2000) at a significantly older level. In New Locality 92*, Beaver Meadow Creek.–Above Angel Falls, York, the boundary is best recognized using conodonts in the town of Java (Wyoming County), a long and inter- (Over, 1997a, b), as commented upon earlier (Text-fig. 16). mittent section is exposed along a tributary to Buffalo Creek Gephuroceratids are last seen just below the boundary in (Pepper & de Witt, 1950: section 6). NY Rte. 78 crosses the Irish Gulf (Loc. 90), as recounted earlier, at approximately creek just below Angel Falls (over the Nunda Sandstone) the level where the Gephurocerataceae become extinct in- and approximately 150 yd (137 m) above the road the upper ternationally. part of the Pipe Creek Black Shale forms a small falls. Bed The Corell’s Point Goniatite Bed has been located at a 92/1 is immediately above the falls on the northern bank. number of localities (House, 1968) and these are indicated On the opposite side of the creek there is a ford crossing to on Text-fig. 17. It is clear that the locality at Corell’s Point a small island and Bed 92/2 is on the island side of the ford. (Loc. 97) was known to James Hall and this could be the 60 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

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 between Lake Erie and Java Village (Wyoming County), with the locality numbers used in the text.

source of the type of Truyolsoceras bicostatum that he de- the outlets of Walker and Corell’s creeks, 250 yd (230 m) scribed in 1843. Cheiloceras is present in collections made in southwest of the former, 2.85 mi (4.6 km) west of Brocton the last century and held in the NYSM but the specimens re- (Chatauqua County) at an altitude of approximately 572 ft mained unrecognized (House, 1962: 276). Despite substan- (174 m). A small point there is formed by a ledge of septar- tial collecting at the new localities. no additional goniatite ian nodules, the Corell’s Point Goniatite Bed, to 0.46 m (18 species have been added as a result of our current work. in) in thickness, overlain by 4.6 m (15 ft) of shale with cal- careous concretions. The fauna occurs in the ledge and as- Localities.–Locality numbers with asterisks have sections il- sociated shale. The goniatites include Cheiloceras (C.) lustrated in Text-fig. 17. The Corell’s Point Goniatite Bed amblylobum, Truyolsoceras bicostatum, and Phoenixites con- (House, 1968: 1066) is almost the only horizon in the centricus, the last a species described by House (1962, 1965) Gowanda Shale known to carry goniatites, but there is a under Tornoceras, but which is now referred to Phoenixites rather lower level known in Walnut Creek (Loc. 99). The (Becker, 1993a). Corell’s Point Goniatite Bed has been traced from the type Although apparently known to James Hall (1843: 246), locality on Lake Erie Shore (Chautauqua County) eastward and probably the source of his Goniatites bicostatus Hall, to Gowanda (Erie-Cattaraugus counties), and faunal and 1843, the first clear reference to the bed seems to be by lithological evidence for the horizon has been found as far Luther (1903: 1025) who referred to concretions on the east as Java (Wyoming County). Tesmer (1954, 1975) gave shore west of Brocton that could refer to this locality. The information on Gowanda localities in Chautauqua and Cat- Clarke and Luther collections of Cheiloceras (C.) amblylobum taraugus counties. The main goniatite localities are as follows dated 1898 are at Albany (NYSM 11239-11242). The redis- (Text-fig. 17): covery of this locality in recent times is due to W. Moran Locality 97*, Corell’s Point.–Corell’s Point lies between whose collection is in Washington, DC (USNM 137665 HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 61

and 137666). This section was also illustrated by Baird & 865 ft (264 m). The sequence of the Gowanda Shale cannot Lash (1990: fig. 4). be measured in detail here, but the distinctive fauna and Locality 98*, Little Canadaway Creek.–Eastward the lithology of the goniatite bed make the assignment clear. Corell’s Point Goniatite Bed crosses Little Canadaway Locality 101, Big Indian Creek.–On Big Indian Creek Creek, 2,200 yd (2 km), north-northwest of the junction of (Cattaraugus County), the Corell’s Point Goniatite Bed Lake Road and NY Rte. 20 at Lamberton (Chautauqua crops out 46 m (50 yds) below a track and ford crossing County) at an elevation of approximately 625 ft (190 m). from Olmstead Farm, approximately halfway between The section lies above the creek crossing of the New York Wardtown Road bridge and Planck (or Plank) Road at an al- State Thruway. A bioturbated 12.7 cm (5 in)-thick siltstone titude of approximately 880 ft (268 m). bed is seen approximately just where exposures commence Locality 102, Little Indian Creek– The locality on Little above the thruway overpass, and the goniatite bed is approx- Indian Creek (Cattaraugus County) lies approximately 64 m imately 3.9 m (13 ft) higher. The calcareous ledge is again (70 yd) downstream from the crossing of Planck Road, at an septarian and of irregular thickness, to 0.38 m (15 in). The altitude of 890 ft (271 m). Loose blocks of the unit are com- same fauna recurs as at Corell’s Point. mon for 30 yd (27 m) farther downstream, from which the At the mouth of Little Canadaway Creek, 1.4 mi (2.3 goniatites mentioned here were collected. km) south-southwest of Van Buren Point on Lake Erie Locality 103, Cattaragus Creek Tributary.–On the Cat- Shore, blocks of the Corell’s Point Goniatite Bed with the taraugus Indian Reservation, the Corell’s Point Goniatite typical fauna can be found. Bed was located just above the Mackinaw Road Bridge Locality 99*, Walnut Creek.–On Walnut Creek, there is crossing of a tributary of Cattaraugus Creek, 3 mi (4.8 km) a long section of the Gowanda Shale (Pepper & de Witt, northwest of Gowanda (Cattaraugus County) at an altitude 1951: section 1). The first horizon bearing goniatites was of approximately 900 ft (274 m). found at a level 0.9 m (3 ft) above the siltstone on which Locality 104, Cattaragus Creek, South Branch.–There the Mixer (Sheridan) Road bridge (Loc. 99/a) is built, on is a long and excellent section of the Gowanda Shale along the northern outskirts of Forestville (Chautauqua County). the South Branch of Cattaraugus Creek, southeast of This level yields crushed Truyolsoceras cf. bicostatum more Gowanda (Cattaraugus County). The Corell’s Point Goni- evolute than the type described by Miller (1938: 80) as atite Bed forms a joint-controlled water-shoot (or flume), 1 Tornoceras (Aulatornoceras) clarkei. The locality of Truyol- mi (1.6 km) south-southeast of Forty Bridge, 675 m (739 soceras? clarkei was given by Miller as near Forestville, New yd) south along the creek from a stream and waterfall York, but it could be that the material he described was that marked on the Gowanda 7 1/2’ Quadrangle Map, at an al- referred to by Harris (1899) as from Walnut Creek, 9.1 m titude of 975 ft (297 m). A good goniatite fauna was col- (30 ft) below the railroad bridge and approximately 50 m lected from this locality. (10 rods or 165 ft) downcreek from it. This is in the upper Locality 105, Clear Creek, North Branch, Taylor Hol- part of the Gowanda, and not the Laona Sandstone or West- low.–Approximately 2.5 mi (4 km) east of Taylor Hollow field Shale, as speculated earlier (House, 1965: 123), but the (Cattauraugus County) is a tributary of the North Branch of material might not be the same. Clear Creek. Approximately 150 yd (140 m) above the The Corell’s Point Goniatite Beds, exposed in Walnut Lenox Road, blocks of the Corell’s Point Goniatite Bed with Creek approximately 300 yd (275 m) upstream from the ?Cheiloceras were located at approximately 970 ft (296 m) al- railroad bridge at Forestville (Loc. 99b), yields Truyolsoceras titude. The associated beds here are siltier than those farther bicostatum and Cheiloceras amblylobum. The elevation of the west. bed is approximately 847 ft (258 m). This locality also seems Locality 106, Clear Creek, North Branch, to have been noted by Harris (1899), who recorded fossils Marshfield.–In Erie County, 2 mi (3.2 km) west-southwest associated with concretions approximately 100 m (20 rods) of Marshfield, on the North Branch of Clear Creek, 100 yd above the railroad culvert. On section 1 of Pepper & de Witt (90 m) downstream from Jenings Road bridge, a soft silt- (1951), the goniatite bed is marked as a layer of concretions stone immediately above a concretionary layer contains 1.5 m (5 ft) above a 0.6 m (2 ft)-thick black shale, 18 m (59 Truyolsoceras cf. bicostatum in a setting similar to that east of ft) below the base of the Laona Sandstone. Taylor Hollow. The altitude is approximately 1,050 ft (320 Locality 100, Smith Mills.–In Chautauqua County, the m). Corell’s Point Goniatite Bed is exposed again approximately Locality 107, Anthony Gulf.–On the eastern side of An- 150 yd (137 m) upstream from the King Road bridge cross- thony Gulf, 2.4 mi (4 km) north-northeast of Boston Cen- ing of a tributary of Silver Creek, approximately 0.25 mi ter (Erie County), Cheiloceras was found in the typical (0.4 km) west of Smith Mills at an altitude of approximately recrystallized preservation of the Corell’s Point Goniatite 62 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

Bed, just below a pipeline crossing at an altitude of 1,350 ft Shale. This is thought to be a Famennian IIB level in terms (411 m). The block was not in situ, but can scarcely have of the German sequence (House, 1962) or UD II-G in more been far displaced. modern terms (Becker & House, 2000). Locality 108, Gears Creek.–On Gears Creek, 1.8 mi (2.9 km) south-southwest of Holland (Erie County), a loose CONEWANGO GROUP block of the Corell’s Point Goniatite Bed with ?Phoenixites No goniatites are known this high in the sequence in New sp. in the usual preservation was found at an altitude of ap- York State but faunas are known in adjacent Erie County, proximately 1,400 ft (427 m). Pennsylvania (Loc. 111, below) and there are three Famenn- Locality 109, Java.–A specimen of Cheiloceras (Cheilo- ian goniatite and clymenid faunas known near Cleveland, ceras) amblylobum in the NYSM is labeled “Portage, Java,” Ohio, in the Cleveland Shale and earliest Bedford Shale presumably referring to a locality near Java (Wyoming (House et al., 1986). According to Oliver et al. (1969), the County). Despite a careful search, the Corell’s Point Goni- base of the Cleveland Shale where the lowest of the three atite Bed was not located in place. H. S. Williams (1887: Cleveland faunas occurs (just above the extreme western- 49), however, gave a section in which he recorded goniatites most tongue of the Chagrin), lies at approximately the level and this deserves further attention. Java is the farthest east of the Wolf Creek and Panama conglomerates of New York that Cheiloceras has been identified. Two specimens of State, and hence all three Cleveland faunas would be referred Phoenexites concentricus (NYSM 11965 and 11966) are to the Conewango Group. No attempt has been made in thought to have come from Gowanda Shale equivalents at this work to trace the Ohio levels eastward into Pennsylvania Java. and New York.

CONNEAUT GROUP Localities.– Only two goniatites are recorded from the Conneaut Group Locality 111, Howard Quarries.–North-northeast of of New York State. The only specimen of these that can be Franklin Corners, 14 mi (22 km) southwest of Erie (Erie satisfactorily determined is USNM 137645, described here County), Pennsylvania, at Howard Quarries, is the type lo- as Maeneceras aff. acutolaterale (G. & F. Sandberger, 1850) cality of Maeneceras milleri (Flower & Caster, 1935). White and formerly assigned to Maeneceras cf. pompeckji (1881: 103) described the quarries here and noted that Falls (Wedekind, 1918) (House & Kirchgasser, 1993: 276). The Run flows northward past the quarry. The horizon was said locality is in the Ellicott Shale at Porter Creek (Loc. 110), to be “probably lower Conewango, near the horizon of the near Summerdale (Chautauga County). The other specimen Panama conglomerate” (Miller, 1938: 176). The lower Ve- (NYSM 4090) is the holotype of Tornoceras edwinhalli nango Sandstone is said to occur just below the quarry out- Clarke (1898: 111, text-fig. 85), which is recorded as from crops. The specimen was determined by House (1962: 262) the Conneaut at Nile (Allegany County). A subsequent ob- as Sporadoceras milleri, but would now be referred to servation by G. H. Chadwick reported by Miller (1938: Maeneceras (following Becker, 1993a) and is so described 152) suggests that the specimen could be from either the here; it is correlated with the Cheiloceras Stufe of Europe Chagrin or Volusia member of the Conneaut; the level is (UD II-H); the level of the lowest of the three Cleveland not satisfactorily determined. The specimen is described faunas, referred to above, is referred to the Platyclymenia below as “family and genus indeterminate.” Stufe (UD IV-A).

Localities.– PALEONTOLOGY Locality 110, Porter Creek, Summerdale.–A locality near This section comprises introductory discussions on the gen- Summerdale (Chautauqua County) was the source of a go- eral problems relating to this work and a review of the local niatite collected by H. S. Williams (his locality 518B; details stratigraphic and ecological setting of the late Devonian go- are in his notebooks preserved at the USGS, Washington, niatite faunas in New York State. A review of the evolution DC). The specimen (USNM 137654) is here referred to of Devonian ammonoids, as it applies to the faunas under Maeneceras aff. acutolaterale. J. W. Wells found the locality description, and an account of the terminology and proce- 518B marked on a map belonging to H. S. Williams. The dures adopted in this work are followed by the systematic locality is 2 mi (3.2 km) “northwest of Summerdale along description of the faunas themselves. The last, which consti- Porter Creek at a point just below Porter Cemetery and at tutes the major section, is intended to review all the faunas an altitude of 1450 feet [442 m]” (House, 1962: 277). On known to us and to illustrate type material photographically the geologic map of Tesmer (1954: 29), the locality falls in for the first time. Only brief descriptions and comments on the northern (middle) part of the outcrop of the Ellicott new additional information are given in the cases of those HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 63

taxa for which good accounts are already available in the lit- PALEOENVIRONMENTS erature. Three sedimentological controls influence the nature of the The prime purpose of this work was to establish the de- late Devonian rocks in upstate New York, and hence the dis- tailed biostratigraphic occurrence of late Devonian am- tribution of goniatites. (1) In space is the dominance of the monoids in New York State. That the New York goniatite thick clastic sequences of the Catskill Delta in the east, and faunas were remarkably diverse and well developed has long associated with it, the regular reduction in thickness, and been known. However, the localities of most of the known finer grade of sediment encountered westward where, in the faunas were in only the crudest way related to each other in earlier parts of the sequence, substantial unconformities can a biostratigraphic sense. Hence the necessity for a detailed be documented (Rickard, 1975; Baird, 1979; Baird & Brett, establishment of the stratigraphical relations of the many 1986a, b). (2) In time is the influence of a series of transgres- sections available for study, and review of these has been sive pulses, commencing with that of the Tully Limestone, given in the previous section. A start is made in this section and later by black shale tongues extending eastward into the to give a more detailed analysis of the available goniatite fau- clastic facies and represented successively by the black shales nas, and to this end general comments on the stratigraphic of the Geneseo, Middlesex, Rhinestreet, Pipe Creek, and occurrence and ecology of the faunas are given. Little infor- Dunkirk members (Text-fig. 2); the onset of four of the last mation of this type is available for Devonian faunas else- are mirrored in the lithostratigraphical nomenclature by where in the world largely because most classical Devonian forming the base of group units (Rickard, 1975). The cycles ammonoids are from pelagic facies. Some evidence will be thus initiated are essentially shallowing-upward sequences. provided suggesting that there is rather more facies control (3) In general, these easterly extending transgressive pulses of goniatite distribution than has generally been conceded, show decreasing effect through time. The first here consid- particularly with regard to the Triainoceratidae. ered, that of the Upper Tully, which initiated the Taghanic The goniatite sequence for Frasnian equivalents that is Onlap, extends the farthest. The later black shale tongues now established in New York is one of the more detailed are progressively more limited to western areas. All of this is known in the world, and enables a more critical look at var- illustrated on Text-fig. 3. Corresponding with this, the clas- ious aspects of goniatite evolution. There is considerable tic facies belts extend farther to the west with time, as has scope for additional work of this type even in New York, been recognized for many years (Rickard, 1964). and substantially more in areas with other good sequences of Goniatite occurences are intimately linked to certain fa- this age, as in Belgium, Germany, North Africa, Western cies belts that change their position, during the time period Australia, and the Timan region of Russia especially. In a considered, in relation to the transgressive pulses. Hence it sense, then, the present study is a start only. There are a large is convenient to give generalizations concerning goniatite number of late Devonian genera that have not been found distribution and paleocology under headings of lithological in New York or elsewhere in North America, and hence not facies types. Later sections will deal with geographical distri- placed in the time sequence here established. It is also true butions and with several distinctive organic associations. in some cases that there are no critical grounds for placing the missing genera into the sequence in other ways using Facies Types correlative methods. It is to be hoped that in the future, con- The major facies belts typical of the sequence from the odont studies of type material and other means will enable Taghanic to Chautauquan are illustrated in Text-fig. 18, closer time documentation of goniatite taxa. which is based in part on Bowen et al. (1974). The diagram Future contributions will depend largely on new collect- is largely based on the Sonyea Group, and the geographical ing. We are conscious that only limited time has been spent limits indicated apply to that group only. extending the geographical range of known levels of goni- atite occurrence. Also there are certain levels that have not Clastic facies.–Most of the clastic facies discussed fit readily been searched systematically, notably the black shales them- into the sedimentological model illustrated in Text-fig. 18. selves. But the data recorded here have been assembled over The carbonate-dominated facies, however, apart from the 40 yr and it seems unlikely that delay in recording the data concretions, generally do not, and each merits separate con- would serve much purpose. Rather, it is hoped, this report sideration in the text. will encourage others to study particular levels or problems (1) Black Shale.–This facies of black to grayish-black or in New York and work toward compiling similarly detailed brown to olive-black shale, often massive and weathering data on Devonian goniatite occurrences in other areas of the with extremes of fissility, characterizes the major and minor world. black shale tongues between the Geneseo Shale Member and the Dunkirk Shale Member (Text-fig. 3). The thicknesses 64 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

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

range from a few millimeters to the great sequence of the readily interpreted as part of a shallow setting. The setting Rhinestreet Shale Member, which reaches over 60 m (200 ft) suggested is one in which the sediment/water interface is at outcrop and is very extensive in the subsurface (Rickard, below the wave agitation zone and sufficiently below the 1975, 1981, 1984). Some documentation of the precise col- thermocline to give euxinic and anaerobic conditions within ors involved was given by de Witt & Colton (1978). Within the muds. Essentially these are infill deposits following deep- the generally black shales, lighter bands can occur, and these ening events. generally enter and develop eastward, splitting up the se- References cited by Brett & Ver Straeten (1997) and quences. As will be apparent from Text-figs 3, 8, 11, 13, 15, Murphy et al. (2000) provide access to the vast literature on and 16, these black shales occur as the initiating facies in the relations of tectonics, sea level. and water depth in the both minor sedimentary rhythms and the large sedimentary deposition of black and gray shales in the Appalachian cycles, or rhythms, which define the groups. Basin. Particularly relevant to this study is the work of Mur- Goniatites are very rare in this facies with the exception phy et al. (2000), a fine-scale biogeochemical analysis of core of a few horizons. Whether this rarity is due to early diage- samples through the facies of the upper Hamilton and lower netic dissolution of the aragonite goniatite shells has not Genesee Groups in New York. They proposed a model in been determined. Levels with crushed impressions generally which the enhanced organic carbon burial in black shale show little evidence of either the shell or of shell replacement units (like the Geneseo Shale) was the result of high surface- mineralization as, for example, in the Geneseo and Middle- water productivity resulting from recycling, from the bot- sex Shales. Exceptionally solid specimens occur either as cal- tom to the surface, of the limiting nutrient phosphorus, a citic replacements and infills in calcareous nodules, as in feed-back mechanism in a shallow marine ecosystem. Their some levels of the Rhinestreet Shale, or as pyritic internal interpretation does not support water-column stratification molds, as rarely in the Angola Shale. Associated benthic (i. e., anoxic conditions under a pycnocline) but rather limits fauna is extremely limited, consisting largely of small lingu- the hypoxic conditions (a critical limiting factor for the ben- loid brachiopods. The shales often appear rich in microflora. thic biota) to levels only at or near the sediment/water inter- Interpretation of this facies is part of the much-discussed face. The elevated productivity would enhance the ecological problem of the American Paleozoic black shale province as opportunities for groups like goniatites that occupied niches a whole. As suggested earlier (House, 1975b: 477), excessive higher in the water column. Following their model, the rar- depths of formation seem unlikely. Sutton (1963: 97) gave ity of goniatites in black shales would largely be the result of depth estimates of probably several hundred feet and possi- taphonomic removal. bly several thousand feet for the formation of the black (2) Gray to Olive Shales and Mudrock.–As with the shales of New York. Lower figures, however, seem preferable black shale facies, there is a considerable range of color rep- for the following reasons: (1) The generally cratonic setting resented in this wide group. Variations of gray are common- of the facies, apart from the Michigan Basin, make depths of est. Olive-green coloration characterizes much of the several thousands of feet difficult to conceive on geophysical Cashaqua Shale facies of the classic Naples type, but also grounds. (2) The increasing evidence of unconformities tends to enter with increasing siltiness. There are transitions within the westerly black shale type regimes (Rickard, 1975; also into the black shale colors. Baird & Brett, 1986a, b; Ettensohn, 1994) indicate eleva- Goniatites are not rare in this facies, and are in places tion and erosion more readily associated with shallowness common. Generally they occur in a characteristic preserva- than extreme depth. (3) The evidence of winnowed and re- tion type with calcitic inner whorls and crushed outer manié levels, such as the North Evans Limestone, is more whorls. Of course, it is in these shales that the goniatite-rich HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 65

concretions occur, but these represent a different seafloor shelled benthos. Burrowing in these siltstones is rare. environment and are treated separately. Nevertheless, the oc- (4) Turbidites.–Some, if not many, of the siltstones dis- currence of solid, calcitic inner whorls probably represents cussed above could be turbiditic in origin, but some are iso- the nascent development of the concretion facies. It is no- lated turbidite beds within shale facies (see 1 and 2 above). ticeable that crushed remains are generally absent. Some turbidite beds have been traced for considerable dis- In the facies transect of Text-fig. 18, bioturbation and tances. evidence of trace fossils enter in with the gray shale facies of At the mouth of Farnham Creek, Lake Erie Shore (Loc. the basin and open shelf. Associated faunas, excluding those 72, Text-fig. 15) is one such turbidite, approximately 20 cm of the concretions, include commonly abundant occurrences (8 in) thick, within the seventh microcyclothem or rhythm of bivalves such as Pterochaenia, rarer Buchiola, and other from the base of the Angola Shale (West Falls Group). This smaller forms. These might well have been epiplanktonic turbidite unit, varying little in thickness, has been traced at (House, 1975b: 482; Grimm, 1998). At certain levels in the the same level for 50 mi (32 km) to the east (occurrences Genesee Group, the top surfaces of dark gray shales toward are shown on Text-fig. 15). This can be referred to as the black have exceedingly abundant accumulations of “Farnham Creek Bed.” No goniatites are associated with it. Leiorhynchus and Barroisella (Thayer, 1974). Ager (1967) A similar occurrence is the Bluff Point Siltstone, a 1.5- supported an epiplanktonic role for Leiorhynchus in this fa- 11 cm (0.6-4.3 in)-thick convolute-bedded unit seen within cies; this interpretation has its origins in the classic essay of the gray shales of the West River Shale (Genesee Group) at Schmidt (1935). Thompson & Newton (1987), however, most localities between the Genesee Valley and Seneca Lake. reinterpreted the Leiorhynchus clusters in the Geneseo Shale It is a level that can be traced farther to the east and west as opportunistic epifaunal assemblages inhabiting restricted (deWitt & Colton, 1978: pls 2-6). The goniatites (Koenen- dysaerobic environments at the pycnocline/slope boundary. ites and Lobotornoceras) occur in the cycle below it. The relative rarity of shelled benthos is still noteworthy. The Crosby Sandstone is a marker bed at the contact be- (3) Fine Siltstones.–Clastic tongues of well-bedded fine- tween the Penn Yan Shale and overlying Ithaca Shale and grained siltstones, usually well cemented, range in color Sandstone in the vinicity of Keuka Lake; there calcareous from light gray to olive; dark colors are unusual. These units concretions in its base contain the goniatites Koenenites and tend to characterize the upper, coarser parts of major Tornoceras. As noted in the stratigraphic section above, the rhythms or cycles and, more unusually, minor rhythms. For Crosby correlates west to Canandaigua Lake to the “hiatus the major rhythms, they suggest progradation in the final concretion horizon” of Baird (1976) in the Upper Penn Yan stages of basin infill when little accumulation space remains. and eastward as a prominent siltstone within the Ithaca to Often substantial thicknesses are involved (Text-fig. 3). Frag- sections in the Seneca and Cayuga valleys (de Witt & mental shell-bearing siltstones occur at some levels and these Colton, 1978: pl. 3). The Crosby is a complex, bioturbated, seem to be facies intermediates approaching that of silty multi-event bed, the probable turbidite origins of which are sandstones and sandstones discussed briefly below in which obscure. rich shelly faunas often occur. In the Namurian of Devon, England, goniatite nuclei In this facies, goniatites are often common but tend to are commonly found with broken shells of large specimens be restricted to the bases of siltstones. Specimens include the at the base of similar turbidite units (Pinkhoe, Brickpit, Ex- largest known of certain genera, notably of mantiococeratids eter). The New York occurrences differ in that the goniatites and triainoceratids, and, particularly in the Gardeau and En- are large and commonly complete. It could be that they rep- field, respectively. Specimens are preserved solid or with resent shells swept down from higher levels on the paleos- some crushing of outer whorls essentially as siltstone infills lope with the onset of a turbidite flow. without calcitic inner whorls. Early diagenetic matrix ce- (5) Silty Sandstones and Sandstones.–In rocks of this fa- mentation is suggested. cies, which present more clastic rocks to the east, goniatites The limited fauna of this facies is mostly associated with are virtually unknown apart from isolated occurrences. This the goniatites and takes the form of shell incrustations often is the facies that embraces the delta platform, delta front, indicating growth on dead shells on the sea floor. Large spec- and pro-delta facies described, for example, by Bowen et al. imens of orbiculoid brachiopods (Clarke, 1899a; 1899b: pl. (1974). A very rich shelly fauna occurs here that is well doc- 3, fig. 4) growing on the inside of large body chambers are umented as the classic Naples Fauna but does not concern known; encountered more frequently are growths of aulo- this discussion. Yet farther landward, alluvial facies yield no poroid corals in the same position within empty body cham- goniatites at all despite the evidence that the shells of the bers. This suggests that this facies is approaching a better living Nautilus are often blown or swept well onshore aerated environment and represents the entry of calcitic- (House, 1987). 66 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

Carbonate Facies.–The Tully and Genundewa represent the the Tully, shows that intermittent seafloor elevation led to only significant limestone tongues in the part of the se- winnowing and erosion in those areas as they had in the quence dealt with here. The latter, because of its distinct na- Hamilton. Baird & Brett (2001, 2003) and Baird, Brett & ture warrants individual comment. Otherwise, carbonate Bartholomew (2003) have differentiated parts of the Tully as occurrences are localized, taking the form of concretions of highstand and lowstand deposits. They recognize a major Knollenkalk and Kramenzelkalk facies. The general absence lowstand event in the Upper Tully marked by a regional dis- of a carbonate facies belt in the later Devonian of New York conformity at the base of the Bellona Bed in western New is largely responsible for giving it such a distinctive overall York and the base of the West Brook Shale in eastern New character, in sedimentological, paleontological, and paleoe- York. This lowstand marks the reestablishment of the cological terms. Hamilton Fauna (brachiopod-coral biofacies); the subse- (1) Limestones.–Only the Tully Limestone truly belongs quent deepening (West Brook Bed-Moravia Bed) is marked here and this has been the subject of detailed treatment in by the incursion of Pharciceras and the highstand deposits of Heckel’s (1973) magnificent monograph and more recently the Fillmore Glen Bed and post-Tully Geneseo Shale. in Baird & Brett (2003) and Baird, Brett & Bartholomew After the Tully Limestone, good limestones are not de- (2003). As has been documented earlier, only one level veloped higher in the Devonian of New York, but a similar within the Moravia Bed or West Brook Member (Cooper interpretation applies to the limited limestones of the Lodi & Williams, 1935) has yielded goniatites. and Genundewa. Both of these are characterized by the The Tully Limestone level has been regarded as trans- abundance of ammonoids (respectively, Ponticeras and Koe- gressive since the late nineteenth century (Frech, 1897: 255), nenites especially) again indicating a link with the world a view confirmed by the work of Cooper (1968), Cooper & oceans. The styliolinid facies of the Genundewa is a pelagic Williams (1935), Cooper et al. (1942), and Heckel (1973, deposit, but the styliolines, thought to be planktonic, could 1997). Formerly it was regarded as the Upper Devonian well have been concentrated by drifting. No great depth transgression, but the name Taghanic Onlap (Johnson, need be required and the vertically orientated goniatites in 1970, 1989) is appropriate because the actions of the Sub- the Lodi (Pl. 4, Fig. 5) suggest relatively shallow water. The commission on Devonian Stratigraphy (SDS) and Interna- interpretation of the Tully, Lodi, and Genundewa limestones tional Union of Geological Sciences (IUGS) have relegated as relatively deep in relation to the clastic facies has been il- the level to the Middle Devonian (Klapper et al., 1987). It lustrated in recent sea-level curves for New York (House & might be expected that the limestone phase of major Kirchgasser, 1993; House, Menner et al., 2000). rhythms would represent the shallowest facies, but it is best (2) Large Concretions.–Characteristically developed at explained by deepening, for the following reasons. (1) In the specific levels within the shale facies are concretionary hori- general paleogeographical setting of the Devonian Ap- zons. The colors of the concretions tend to match the colors palachian Gulf, clastic facies predominate and the Tully of the shales in which they occur. The larger concretions, Limestones will represent a period when clastic deposition taken arbitrarily as those over 20 cm in diameter, are consid- was far distant; this is most easily achieved by sea-level rise. ered here. Usually these are depressed-ovoid in vertical sec- (2) The entry of “exotic” elements, such as harpid trilobites, tion and circular in plan. Some larger examples show the brachiopod Hypothyridina, and the goniatite Pharciceras, interpenetration of sphaeroids. Concretions are usually sep- indicate links to world oceans and ready migration, such as arated by distances in excess of their width along specific is best explicable by deepening. (It should be noted that Hy- horizons. Rarely they coalesce as concretionary limestones, pothyridina venustula (Hall, 1867) was assigned to a new of which the Corell’s Point Goniatite Bed (Loc. 97) is the genus, Tullypothyridina, by Sartenaer in 2003.) (3) In the best example, which can be traced inland from the shore of New York facies transect, as demonstrated by Cooper & Lake Erie. Some of the more spherical concretions in the Williams (1935: text-fig. 3), Johnson & Friedman (1969), Genesee Group reach over a meter in diameter, and more Rickard (1975, 1981), Baird & Brett (2003), and Baird, depressed examples in the Rhinestreet Shale near Lake Erie Brett & Bartholomew (2003), the Tully facies tongues far reach more than twice that size. to the east, carrying marine neritic facies within successions The largest concretionary levels in minor rhythms or cy- that also bear the Gilboa Forest in a way most simply ex- cles (Text-fig. 19C) tend to occur in the upper part of the plained by transgression. That is not to suggest that great rhythm or cycle, below the black shale that introduces the water depths are required to explain the Tully Limestone succeeding unit. In general, there is also a similar relation in faunas, but transgression clearly led to the facies change. the major rhythms or cycles. For example, larger type con- Westward, the thinning of rock units, and the evidence of cretions occur in the upper part of the West River Shale and unconformities (Text-fig. 7) both below, within, and after lower Cashaqua Shale on either side of the intervening Mid- HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 67

dlesex black shale. Other large concretions occur within the within minor rhythms (Text-fig. 19C); these can rarely be uppermost part of the Cashaqua Shale and upper septarian. Their color is similar to the shales in which they Rhinestreet Shale, on either side of the main Rhinestreet occur. Generally, with decreasing diameter, there is a trend black shale tongue. The largest concretions of the Hanover to greater sphericity. Shale occur a little above the black shale tongue of the Pipe The smallest concretions often show a pyritic nucleus Creek Shale. As will be discussed later, for the major and evidence that a burrow or other organic center occurs rhythms, this introduces an element of cyclicity into the pat- within them. In places, as not uncommonly in the Cashaqua tern. Shale, the nodule can comprise a goniatite body chamber, Concretions tend more frequently to be septarian where but usually not the phragmocone. In smaller concretions, the surrounding shale is darker gray or blacker in color. At evidence of nucleation of the concretion around central or- certain levels, and perhaps areas, the normal crystalline cal- ganic material is more often seen than in larger concretions, citic infill has been replaced by barite, and fossil shells have even those containing goniatites. also been so replaced. The exceptionally fine development of This group of concretions appears to have formed in this type is seen in the Shurtleff Septarian Horizon (Text- progressively deeper and less calcareous sedimentary envi- fig. 10); other levels will be referred to in the section on ronments than the large concretions, but this is an overall preservation. The goniatite beds we refer to as “Fossil Log impression and not universally tenable. Where goniatites are Horizons” (FL) are concretionized rafts of driftwood in preserved, they are usually solid and indicate lithification which the plants and shells accumulated as hydrodynami- before any appreciable compaction had taken place. cally similar, lightweight materials as suggested by Maeda & As with the larger concretions, molds of goniatites with Seilacher (1997). recrystallized shells are sometimes replaced by barite or, Fine, solid, goniatite specimens, often difficult to extract, more rarely, silica. Examples of this type of small nodules are common in many of the large concretionary horizons, al- with diagenetic barite rosettes occur in the lower Hanover though perhaps rarest in the largest. Commonly there is an Shale (Pepper et al., 1985). This replacement will have taken associated fauna of gastropods and bivalves, but these are place long after concretion formation. even more difficult to extract, being smaller and mostly re- (4) Knollenkalk and Kramenzelkalk Levels.–Deeper water crystallized replacements. At baritized levels, where acid dis- Devonian deposits of the European Hercynian Facies com- solution enables easy preparation, a great variety of forms monly comprise micritic nodular limestones and cephalo- are known, as described by Clarke (1899a, b) from the pod limestones known as Cephalopodenkalk, Knollenkalk, or Shurtleff Septarian Horizon (Loc. 41). It is interesting that griotte in the European literature. This represents a facies in where concretions start to coalesce, forming essentially a which goniatites occur abundantly at appropriate levels. In concretionary limestone bed, as the Genundewa Limestone places, the more blotchily colored nodular limestones have and Corell’s Point Goniatite Bed, gastropod and bivalve fau- a distinct “ant-eaten” appearance in weathered outcrops. nas increase both in size and abundance. The Genundewa This gave rise to the term Kramenzelkalk for a distinctive fa- Limestone is distinctive in other respects, however, and is cies in which the carbonate nodules and enveloping shales discussed separately below. In the Corell’s Point Goniatite are differentially weathered with the nodules hollowed to Bed, the numerous gastropods are often similar in size to, form a reticulate network of cells. Tucker (1973) considered and roughly homeomorphic with, the associated Cheiloceras. these facies types as pelagic limestones, a term more geneti- Because shells in these concretions are uncrushed, it is cally precise than the facts warrant. apparent they represent early diagenetic cementation events In New York, at certain levels in the Frasnian, nodular perhaps arising from nucleation centers associated with or- limestones of a similar sort occur. These are commonly red ganic remains. The availability of calcium carbonate for the and green in coloration, form distinct if not thick beds, and process, however, indicates that the concretionary levels rep- show a curious weathering. Generally they are coarser in resent different environmental conditions than that of the grain size and more impure than typical European types, background shale above and below. The baritization is even if their characteristics bring them within the termino- thought to be a much later epigenetic event. logical range of European usage. (3) Small Concretions.–There is a series in size of con- This facies seems to belong to a shallower environment cretions from those just considered down to small concre- than those so far discussed, but which is unusual in being as- tions that are essentially nodules approximately 10 mm in sociated with rises on the sea floor. The European examples diameter. Levels with very small concretions are rare. The characterize the crests of substantial seafloor elevations or larger among this group are similar in other characters to Schwellen. The New York examples suggest a similar, if less those already described, and occur in a similar position spectacular, setting. 68 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

The Parrish Limestone, within the Cashaqua Shale been suggested (House, 1975b) that these might have been (Text-fig. 10, Beds 47/2 and 48/1), occurs on the crest of a planktonic bubble-snails, associates in the ecology of the liv- clastic wedge termed the Rock Stream Siltstone, and appears ing Styliola. Goniatites occur abundantly at certain levels in to owe its characteristics to the oddly raised environment the Genundewa, as at Bethany Center (Loc. 24b), together presented by the crest of the wedge. The Parrish Limestone, with bactritids and orthocones in a manner that suggests described by Clarke (1885: 39) as a Kramenzelkalk facies, is drift accumulation. But there is also a substantial benthonic a famous source of manticoceratids in the region close to fauna (Clarke, 1904; Sass, 1951) that probably flourished Naples (Clarke, 1885; Kirchgasser 1965, 1975). Passing lat- in the accumulating styliolinid sands. erally from this region, both to the west and to the east, the Styliolinid limestones with goniatites are developed to facies changes to shales with nodules such as those previously thicknesses reaching 50 mm at levels in the late Penn Yan discussed. Shale, and West River Shale and correlative horizons, in- Two other examples occur higher in the sequence (Text- cluding the Genundewa, have been found in the Harrell fig. 3): levels in Angola Shale equivalents near Varysburg Shale in the region near Landes Post Office, West Virginia (Wyoming County; Text-fig. 15, Loc. 79/10), and on John- (House, 1978), and in the Harrell Shale in central Pennsyl- son Creek (Wyoming County; Loc. 81/10), which are thin vania (Kirchgasser, 1996b). The Squaw Bay Limestone in developments of this type of facies. At both localities there Michigan is a similar facies. Such widespread styliolinid are other levels rather higher but well below the horizons of units with goniatites are interpreted as trangressive phases. “The Trinity” black shales. A notable one with goniatites oc- Styliolinid limestones or concretions, often with goniatites, curs within the West Hill/Nunda interval at Varysburg occur at several levels in the Sonyea Group and West Falls (Gassman Road) (Loc 80/14). Other levels occur in the Group. Dacryoconarids (sensu Fisher, 1962, and Linde- lower part of the Hanover Shale, as in Walnut Creek (Chau- mann, 2002) range into the upper Hanover Shale, and ho- tauqua County; Text-fig. 16, Loc. 89a), and red and green moctenids have been found at the top of the Hanover Shale Kramenzel-type nodular horizons are found in the upper and in the lower Dunkirk Shale (Yochelson & Kirchgasser, Hanover Shale in Beaver Meadow Creek, Java (Loc. 92). 1986; Over, 1997a, b; Over et al., 1997). Also, as it passes eastward, the Corell’s Point Goniatite Bed appears to lose its concretionary nature, and specimens of MODE OF GONIATITE PRESERVATION typical Cheiloceras from the Java region (Wyoming County) Goniatites in the New York Devonian can be preserved in a appear to have come from nodular limestones; it could be wide range of ways but a few types are typical. These are that the facies recurs higher in equivalents of the Gowanda largely a result of the variety of facies types in which they Shale. occur and hence reflect the classification of facies already re- (5) Styliolinid Limestones.–Quite an unusual and sepa- viewed. rate facies is represented by calcareous levels where the small No goniatites preserving the original aragonite shell ma- conical shell of Styliolina fissurella (Hall, 1845), and perhaps terial are known in the New York Devonian. The shells, if other dacryoconarids, become so important as to form dis- preserved, are invariably recrystallized, normally as calcite tinct units. The best developed level of this sort is the Ge- but also, rarely, as barite, as with the material of the Shurtleff nundewa Limestone, but other levels of a similar sort occur Septarian Horizon (Loc. 41/1) in the upper Cashaqua Shale in the Penn Yan Shale below and the West River Shale above. in Livingston County. Larger specimens usually lose even Recognition of the Genundewa Limestone by styliolinid oc- this, and such shells are represented by external and internal currence alone is most untrustworthy. A black shale beneath molds. Attention should be drawn to the fact that the exqui- it (Text-fig. 8) is used here as the most accurate means of site material of the inner whorls of goniatites, especially that defining its base. figured by Clarke (1899a, b) and refigured here (for example Whereas the preceding carbonate facies seem to be re- on Pl. 21) was exposed by Clarke by dissolving away recrys- lated to particular depth and site-related factors, the Styli- tallized shells set in mudrock and, when dry, making gutta olina facies relate to a peculiar abundance at particular times percha molds. The resultant preservation is incredibly fine of an organism that is thought to be planktonic and usually but the original specimens were not retrieved. pelagic and whose remains fell to the sea floor after death The commonest of the natural internal molds are those (Fisher, 1962; Lindemann, 2002). Thus their accumulation preserved in pyrite. These rarely exceed 10 mm in diameter probably depended on factors of abundance, wind direction, and are associated with dark gray or black shales, as in the tidal patterns, current movements, and other factors rather West River Shale and Rhinestreet Shale. Remanié accumu- than on seafloor characteristics. Commonly associated with lations of pyritic material, as in the Leicester Pyrite (Text-fig. the styliolinids in the Genundewa are small snails, and it has 7) and other levels are concentrates resulting from the win- HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 69

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 major rhythm or cyclothem; Middlesex Shale at base. B. Diagram simplifying data on facies within the lower West Falls Group of western New York indicating the sequence in another type of major rhythm or cyclothem; Rhinestreet Shale at base. C. Diagram indicating the sequence in a typical minor rhythm or microcyclothem based on examples within the Angola Shale of the West Falls Group. nowing and erosion of shales which formerly contained the Possible causes of such rhythmicity of cyclicity have been pyritic molds (Baird & Brett, 1986a). debated since they were recognized as features of the Jurassic rhythms in the South of England by Conybeare & Phillips CAUSES OF MAJOR AND MINOR RHYTHMS AND CYCLES (1822). For the smaller scale rhythms, usually of the order As has been illustrated (Text-figs 3, 8, 13, etc.), transgressive of 0.5-2.5 m, a cause from climatic changes would depend pulses, especially those initiated by black shales, either at a on changes in outer atmosphere insolation as controlled by major or minor rhythmic scale, form a characteristic feature the changing orbits due to perihelion, precession, and the of the stratigraphy of the late Devonian in New York State. short and longer eccentricity cycles. The resultant climatic Most are clearly of a rhythmic type, with an asymmetrical shifts could cause small sea-level changes due to simple ex- pattern ABCABC or abcabc (Text-figs 19A, C). As noted pansion, the freezing and melting of polar and mountain above, Murphy et al. (2000) have proposed a within-basin ice; on land such changes of climate would modify vegeta- recycling model in interpreting the geochemistry of black tion cover, rates of weathering and erosion, and the nature and gray shale cycles in the upper Hamilton and lower of sediment reaching the basin. The New York sequence Genesee Groups. As has been deduced from previous con- with which we are concerned shows no long-continued sideration, the ABC, abc sequences are interpreted as shal- small-scale rhythmicity as does, for example, the Devonian lowing upward sequences with the shallowest facies (C) pelagic facies of the Montagne Noire (House, 1995). Small- abruptly followed by deepest facies (A). In contrast, the scale (meter-scale) rhymicity occurs in parts of the New York major cycle of the Sonyea Group with a pattern ABCBA succession, most notably in the West River Shale (the (Text-fig. 19A) is a true symmetrical cycle. In this case, the twenty-three rhythms) and in the lower Angola and lower shallowest facies (C) is succeeded (gradually) by deeper water Hanover shales as discussed in the stratigraphy section. Nev- facies (B) which in turn is followed by the deepest facies (A). ertheless, there are too many transgressive pulses perhaps 70 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

due to other causes, such as tectonics, which interrupt sed- approximately corresponds to the lower Assise de Frasnes imentation, to make the New York succession easy for se- onlap of Europe. Again the contrast between the lower and quence or cyclostratigraphic analysis. Primary evidence for upper parts and the preceding black shale indicate that this the operation of Milankovitch cycles is therefore unlikely to is also a polyphased event. It corresponds to depophase IIb be forthcoming, and interpretation must be by analogy with of Johnson et al. (1985). those parts of the stratigraphical column now well docu- The Middlesex black shale pulse seems to correspond mented in Milankovitch and oribital forcing terms. with the deepening that resulted in the growth of Belgian In the New York setting, interpretations of larger scale F2d reefs (House, 1985). The same level appears to be rec- rhythms fall into two main groups. First are those invoking ognizable as the base of the Domanik in northern European intermittent basinal subsidence probably caused by periodic Russia (House, Menner et al., 2000) and the same pulse of movement on faults bounding depositional basins associated deepening has been recognized in Western Australia (Becker with convergent-margin tectonism (the tectophases of Et- et al., 1993). This is the base of depophase IIc in the termi- tensohn, 1985, 1994). Second are those invoking global eu- nology of Johnson et al. (1985). static rises of sea level (see references below). Theories of the The major deepening event represented by the latter group would accept the importance of differential Rhinestreet black shale can similarly be correlated in other basinal subsidence in explaining the pattern of thickness areas, for example with the initiation of the F2h reefs in the variations in the New York Devonian, but would consider Ardennes (House, 1985). This appears to be the base of de- the subsidence to be generally gradual and not intermittent. pophase IId in the terminology of Johnson et al. (1985) The discrimination between these groups of hypotheses whose major depophase boundaries were linked to the group really depends on whether transgressive pulses can be corre- terminology of Rickard (1964, 1975), in this case the base lated over wide areas of the globe in Devonian time. It is of the West Falls Group. But some (Sandberg et al., 2002) thought that for the group of strata with which we are con- seem to be taking the base at the later semichatovae trans- cerned there is increasing evidence that this can be sustained, gression, which could correspond to one of the later trans- and that the second group of hypotheses seems generally to gressive pulses in the Rhinestreet recognized by Sutton have been the more important. Increasing evidence in favor (1960, 1963), but this interval is still poorly documented of the latter group of hypotheses includes the observation by conodonts in New York (Klapper et al., 1995). that many of the major facies movements are international The Pipe Creek black shale is a widely recognized inter- in scope (House, 1983, 1985; Johnson et al., 1985, 1986), national transgressive event (dysoxic pulse) that corresponds but especially the detailed correlations of the Frasnian in approximately to the German Lower Kellwasserkalk trans- Western Australia (Becker et al., 1993), northern European gression and the initiation of the deep-water Schistes de Russia (House, Menner et al., 2000), and New York (House Matagne in Belgium (Schindler, 1990, 1993; House, 1985, & Kirchgasser, 1993) 2002). No zonable conodonts are known from the Pipe The main transgression indicated by the Tully Limestone Creek but the level is probably within Zone MN12 (?Upper and especially the West Brook Member or Upper Tully di- rhenana) (Over, 1997a, b; Klapper & Becker, 1999). The visions (Heckel, 1973; Baird & Brett, 2001, 2003; Baird, Kellwasserkalk transgressions in Europe were first discussed Brett & Bartholomew, 2003) has been widely recognized in in detail by Buggisch (1972) and recognition was then ex- North America as the Taghanic Onlap by Johnson (1970), tended to North Africa by Buggisch & Clausen (1972). The which is just a new name for the American Upper Devonian correlation of the Pipe Creek Shale with the Lower Kell- transgression of the consistent literature of earlier scientists wasserkalk of Germany was not recognized by Johnson et al. dating back a century. Internationally, the importance of (1985, 1986), who correlated the base of the Rhinestreet onlap movements at this time-interval have been docu- Shale with the Lower Kellwasserkalk, a view now known to mented by House (1975a). It is now recognized, however, be in error. that in New York, several phases of transgression are in- Through the work of Over (1997a, b, 2002; Over et al., volved (Text-fig. 7) and much has still to be learned approx- 1997), much more detailed correlation is now available for imating these, both with regard to sea-level changes and the placing of the newly defined Frasnian/Famennian extinctions, internationally as well as locally in New York. boundary (Klapper et al., 1987). It clearly precedes the base Depophase IIa of Johnson et al. (1985) commences at the of the major tongue of the Dunkirk Shale (boundary base of the Tully Formation. marked in Text-fig. 16). Thus the underlying black shale, The Genundewa has also been interpreted as a transgres- known informally by John Huddle as the Rider Bed (from sive level in New York (House, 1983, 1985; House & Kirch- the miner’s term for any underlying or overlying leaf from a gasser, 1993; House, Menner et al., 2000). The level coal seam), seems to be the New York equivalent of at least HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 71

part of the Upper Kellwasserkalk. In view of the detailed doc- 2002). umentation of the phases of the Kellwasser Events (Schindler, 1990, 1993), work is still needed to give precise EVOLUTION OF LATE DEVONIAN GONIATITES documentation with Europe and other areas (House, 2002). A current view of the origin and radiation of Devonian go- The Dunkirk black shale pulse corresponds to the Ne- niatites and their Late Devonian derivatives, the clymeniids, hden transgression of Germany and other areas that initiate is illustrated in Text-fig. 20. This also shows the “events,” Cheiloceras Stufe faunas, which was named the Nehden usually marked by widespread international transgression Event by House (1985), but Cheiloceras arrives later in New and anoxia, which apparently had a major role in controlling York. Although the Dunkirk Shale corresponds to the Ne- evolution of the group (House, 1985, 1996; Becker, 1993b; hden Event (Becker, 1993a) and transgression of the early Becker & House, 2000). The group was derived from or- Famennian of Europe, relevant goniatite faunas do not ap- thoconic Bactritida and the coiled goniatites with a dorsal si- pear until the Gowanda Shale. The Dunkirk corresponds to phuncle first appear a little above the base of the Emsian. the base of depophase IIe in the terminology of Johnson et By the beginning of the late Givetian (formerly basal Upper al. (1985, 1986). Devonian), at the Taghanic Event, most earlier Devonian These main events are interpreted as transgressions and stocks had become extinct, including the families Agoniati- most are associated with a flooding of dysoxic- to anoxic- tidae, Pinacitidae, Sobolewiidae, and almost all the Maenio- type sediments. It is not to be expected that the sedimento- ceratidae. logical effects of transgressions will be exactly Formerly, all the Frasnian gonitiates apart from the An- contemporaneous in different areas and where those areas arcestidae and Tornoceratidae were grouped together as the can be related to different positions on the hypsometric Gephuroceratoidea. It is now known, however, that the curve. Pharciceratidae and its derivatives (Eobeloceratidae, Pettero- The increasing transgressions through the Frasnian have ceratidae, Triainoceratidae, and the problematic Devono- been argued as the main cause of late Frasnian extinctions, pronoritidae) have their origin in the Maenioceratidae by and indeed, these are mostly accomplished by the Kell- the proliferation of umbilical lobes (Bensaïd, 1974). wasserkalk transgressions (House, 1975b). It is not thought Unless otherwise noted, the framework of classification that offlap and regression leading to “perched faunas” (John- adopted here corresponds to that adopted generally in recent son, 1974) are a reasonable interpretation because the main years (House, 1981a; Becker & House, 1993, 1994a, b; recognizable regressive events, in Europe as in North Amer- Becker, 1993b). ica, do not appear until the late Cheiloceras Stufe, well after Ranges of genera given here are based on international the extinctions. work resulting from investigations since 1954. More recent Although major global eustatic changes of sea level can are the detailed investigations resulting from work with the explain similar major rhythmic patterns in different conti- Subcommission on Devonian Stratigraphy in the definition nents, a similar interpretation for microcycles cannot be of stage and series boundaries and in relation to those tested because correlative tools are too imprecise. The low “events” that appear to have significantly affected the evolu- order of thickness involved in the minor patterns (Text-fig. tion of the group. The most significant ones are marked in 19C) could be equally well explained in sedimentological Text-fig. 20. This work has been greatly aided by the precise ways alone by migrating distributaries and other mecha- conodont biostratigraphy established for the late Devonian nisms against a general background of basinal and basin- (Ziegler, 1962, 1971; Ziegler & Sandberg, 1990; Klapper, marginal subsidence. 1989, 1997, 2000; Klapper & Becker, 1999) that has given If the global eustatic cause for the major rhythms comes evidence of age independent from the goniatites themselves. to be accepted, then the question of a mechanism for sea- Text-fig. 21 is a range diagram for late Devonian taxa known level changes is required. Two of the main contenders are worldwide over the period with which we are concerned in changes in the amount of polar ice and volumetric changes New York – the late Givetian to the mid-Famenian. Many at constructional plate margins. The glacio-eustatic inter- of these genera are not known in New York or in North pretation is difficult to sustain in the absence of clear evi- America. dence for significant polar ice in the Devonian and the generally warm and equable climate generally inferred in a BIOSTRATIGRAPHY period that represents the maximum in faunal and floral di- versity in the Paleozoic. Other contenders for the sea-level INTERNATIONAL GONIATITE ZONATION changes include orbitally-forced climate changes (House, OF THE LATE DEVONIAN 1985; 2002) and comet-shower impacts (Sandberg et al., Detailed work on the ammonoid zonation of the Devonian 72 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

Text-fig. 20. Diagram illustrating the evolution of Devonian ammonoid families. The width of bars corresponds to the number of genera at particular times as indicated by the scale. The timescale used is based on Becker & House (1994a) and Becker (1993b). Also indicated are the named environmental stress events. Modified from House (1996). HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 73

Text-fig. 21. Diagram illustrating the ranges of late Devonian goniatite genera internationally. From Becker & House (1994b); Famennian data from Becker (1993b). 74 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

essentially began in Germany when Devonian studies fol- UD III Prolobites Stufe lowed the celebrated work of Albert Oppel (1831-1865) on UD II Cheiloceras Stufe the Jurassic. Fritz Frech (1861-1917), August Denckmann Frasnian (1860-1925), Emmanuel Kayser (1845-1927), Rudolf UD I Manticoceras Stufe Wedekind (1881-1961), and Hans Matern (1903-1933) Middle Devonian (MD) were chief among early workers. In more recent times no- Givetian table contributions on the Famennian were made by H. MD III Pharciceras Stufe Schmidt & O. H. Schindewolf, B. I. Bogoslovsky, D. Weyer, MD II Maenioceras Stufe J. D. Price, D. Korn, and R. T. Becker. Eifelian As noted by Becker & House (2000: 115), the potential MD I part Anarcestes Stufe and Pinacites of ammonoids for international correlation within the De- Stufe vonian is limited by the demonstration in recent decades Lower Devonian (LD) that synchronous but widely separated sedimentary basins Emsian can contain different sequences despite the pelagic lifestyle LD III Anetoceras Stufe and part Anarcestes of the group. The faunal distinctions appear to be the result Stufe of many factors including the individual structural and facies Pragian (LD II) No ammonoids known. histories of basins, the limitations of faunal exchange be- Lochkovian (LD I) No ammonoids known. tween open-marine environments by paleogeographical bar- riers resulting in endemic taxa, sedimentary discontinuities, For the Upper Devonian, the former Stufen or major di- and discontinuous representation of ammonoid biofacies. visions or zones of Wedekind & Schindewolf are essentially Nevertheless, using the wealth of new data generated by am- retained, each designated by Roman numerals (UD I-VI) monoid workers from many sections around the world, and the name of a genus. Although never explictly defined, Becker & House (2000) proposed a new and much refined the Stufen are essentially taxon range zones that are based on zonation that is correlated with the celebrated conodont the first and last occurrence of a single taxon. Wedekind & zonation and with the established series and stage bound- Schindewolf subdivided the Stufen using Greek letters; thus aries. UD II, the Cheiloceras Stufe of the Famennian Stage, was Because the major early contributions on late Devonian divided into IIa and IIb. Recent work has added consider- ammonoid biostratigraphy were based on sequences in Ger- able precision to the zonal subdivision of the Stufen. For the many, it is appropriate to review that which has been termed Lower and Middle Devonian, the revised zonation uses the “standard” succession established by the work of Frech, Roman numerals and the Roman alphabet to subdivide the Denckmann, Wedekind, Schindewolf, Matern, Schmidt, Stufen (e. g., MD III-A through E for the Pharciceras Stufe Lang, and others in the Rheinish Schiefergebirge (Rhenish of the upper Givetian Stage) (Becker & House, 1994b). Slate Mountains). A brief account of the characteristics and For the Frasnian Stage of the Upper Devonian, the re- geographical distribution of Devonian ammonoid zones and vised zonation, based largely on ammonoid successions in faunas has been published (House, 1979) and need not be the Canning Basin, Western Australia, and New York, fol- repeated here. It is important to bring this up to date by lows the same methodology. The Manticoceras Stufe, UD I, noting especially the elevation of the lunulicosta Zone (for- is divided into twelve lettered divisions, UD I-A through L, merly included in the Manticoceras Stufe) to a new Pharci- which replace the four divisions (I alpha to I delta) of ceras Stufe (House, 1985) in the position below the now Wedekind. The new subdivisions for the Upper Devonian as much-raised conodont-defined Middle/Upper Devonian a whole are shown in Text-figs 20-21 and for the Frasnian boundary (Klapper et al., 1987). The discussion that follows and Famennian in Text-fig. 22. is a brief introduction to the data, methodology, and termi- The divisions in the zonation of Becker & House (2000) nology of the international ammonoid zonation of Becker & are essentially interval zones, defined primarily by the entry House (2000). of genera rather than the occurrence of specifically named The major subdivisions of Devonian ammonoid zona- zonal species. The entry of genera is more appropriate for tion are: international use because each area that has been examined Upper Devonian (UD) (New York included) has a regional zonation, the character- Famennian istic feature of which is the dominance of local (often en- UD VI Wocklumeria Stufe demic) species or subspecies. The new international UD V Clymenia Stufe divisions are thus defined as Ammonoid Genozones. Becker UD IV Platyclymenia Stufe & House (2000) also introduced an idealized parallel HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 75

Text-fig. 22. Zonation of the late Devonian using ammonoids plotted against the conodont zones. Based on Becker & House (2000). 76 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

scheme of Ammonoid Standard Zones, based on the entry ian groups of the Ammonoidea (Text-fig. 20). of species (zone fossils) with international or interbasinal distribution but that might not occur in all basins with con- Pharciceras Stufe, MD III (A–E).–The succeeding lunuli- temporaneous ammonoid faunas. For regional successions costa Zone (I alpha) of Wedekind (now Pharciceras Stufe, with less cosmopolitan or endemic forms, Becker & House MD III) was first defined by Frech (1887: 433) using faunas (2000: 115) recognized Ammonoid Regional Zones, which from several localities in the Rhenish Schiefergebirge and are useful for intrabasinal correlation but whose marker Harz Mountains of Germany. The Pharciceras Stufe intro- forms (genera or species) “…may differ from standard or in- duces members of the old suborder Gephuroceratina, first of ternational zones (for example endemics may be used) or the Pharciceratoidea and of the Gephuroceratoidea shortly may have shorter ranges than known from the total global thereafter. Because these groups are thought to be independ- record (regional topozones); for example, delayed appear- ently derived from the Anarcestidae, a single suborder for ance of Manticoceras in various areas in comparison to its them is inappropriate. early appearance in New York.” The broad key to the new The new subdivision of the Pharciceras Stufe recognizes zonation (Text-figs 20-21) shows the international range of five divisions (Becker & House, 1994b, 2000). The fauna of goniatite families and genera in the Frasnian as currently un- MD III-A is defined by the entry of Pharciceras and Mzer- derstood. The zonation of the succeeding Famennian is not rebites but with the hangover from the Maenioceras Stufe of indicated because clymeniids, which form the basis of the Maenioceras, Afromaenioceras, the anarcestid Atlantoceras, subdivision, have yet to be found in New York State. and tornoceratids, especially Tornoceras and perhaps Epi- tornoceras. MD III-B is defined by the entry of multilobed Givetian pharciceratids, especially Stenopharciceras [e. g., S. lunulicosta The faunas of the Eifelian and earliest Givetian of New York (G. & F. Sandberger, 1850)]. MD III-C shows the entry of are not of concern here. A review of known records was Synpharciceras and new eobeloceratids. MD III-D shows the given by House (1962, 1981b) and Becker & House entry of other new eobeloceratids and the earliest acantho- (1994b), but there is important new goniatite information clymeniid, Pseudoprobeloceras. MD III-E shows the entry of in the unpublished thesis of G. Kloc (1983) and in his pri- the Petteroceratidae with Petteroceras, Meropharciceras, and vate collections. the acanthoclymeniid Ponticeras. By its close, all of the eo- beloceratids have become extinct as have pharciceratids apart Maenioceras Stufe, MD II (A-D).–As has already been de- from Neopharciceras and one species of Petteroceras. scribed in the section on the evolution of late Devonian go- niatites, the major break in the ammonoid sequence is at Frasnian the Maenioceras Stufe/Pharciceras Stufe boundary, which was formerly taken in Germany as the Middle/Upper Devonian Manticoceras Stufe, UD I (A–L).–This division corresponds boundary, and by most as the Givetian/Frasnian boundary rather exactly with the Frasnian as now defined. It corre- (House, Feist & Korn, 2000). Toward the end of the Mae- sponds to the dominance of the Gephuroceratidae and Be- nioceras Stufe interval, a wide range of typical Middle De- loceratidae. Formerly it was divided into I beta, I gamma, vonian goniatites become extinct, namely Sobolewia, and I delta by Wedekind (1913, 1917, 1918). Matern Pseudofoordites, Wedekindella, Foordites, Holzapfeloceras, (1931a) combined the two middle zones as I beta (gamma). Cabrieroceras, Werneroceras, Agoniatites, and Sellagoniatites. But work on far more detailed sections than were available All of these genera, except perhaps Foordites, Pseudofoordites, to those authors has led to the new subdivision into 12 di- and Sellagoniatites, the precise ranges of which are uncertain, visions, UD I A-L, the distinction of which is based on work are documented in late M. terebratum Zone faunas (MD II- in Australia (Becker et al., 1993), Russia (Becker et al., 2000; C). For example, most of these genera occur in the Penton- House, Menner et al., 2000), North Africa (Becker & warra Goniatite Bed of North Cornwall (House, 1963). By House, 2000a, b), Germany (House & Ziegler, 1977; the boundary, therefore, the families Pinacitidae, Holzapfe- Becker & House, 1993, 1994a), France (House et al., 1985; loceratidae, and Agoniatitidae become extinct, and the Mae- House, Becker et al., 2000; House, Feist & Korn, 2000), nioceratidae survive for only a very short time. This MD and North America (House, 1962; House & Pedder, 1963; II-D extinction is the Taghanic Event of House (1985). Kirchgasser, 1974, 1975, 1982, 1985; Kirchgasser & House, Only a remnant of the Anarcestidae survives, perhaps to the 1981; House & Kirchgasser, 1993, and herein). early Famennian, and gives rise to the Gephuroceratoidea The fauna of UD I-A, or the Ponticeras Genozone or Di- and Pharciceratoidea. The Tornoceratoidea survived the vision, is subdivisible into three phases, not all of which are Taghanic Event and appear to have given rise to all Famenn- recognized internationally. At the very base, as shown in the HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 77

basal Frasnian GSSP (Global Boundary Stratotype Sections divisions corresponded to the local German Famennian and Points) at Puech de la Suque, Montagne Noire, south- stage-type divisions. Thus, Division II, the Cheiloceras Stufe, ern France (Klapper et al., 1987; House, Feist & Korn, equated with the Nehden; Division III, the Prolobites Stufe, 2000), the record indicates that Peterroceras and Neopharci- and Division IV, the Platyclymenia Stufe, equated with the ceras, the last remnants of the Pharciceratidae, just cross the Hemberg; Division V, the Clymenia Stufe, equated with the boundary; their extinction early in UD I-A marks the Dasberg; and Division VI, the Wocklemeria Stufe, equated Frasnes Event (House, 1985). Then the long-ranging Pon- with the Wocklum. But these stage names are in the process ticeras and Epitornoceras continue with the entry of Chuto- of being redefined in Germany and it is to be expected that ceras marking the base of an upper phase. UD I-B, or the this simple scheme, with its priority, is likely to be replaced. Koenenites Genozone, is marked by the entry of Koenenites The terms were originally lithostratigraphic ones, and con- and the Koenenitidae. UD I-C, the Timanites Genozone, version to biostratigraphic terms, or chronostratigraphic if corresponds to the curious international occurrence of that accepted by the International Union of Geological Sciences genus in the Timan Basin, Russia, western Canada, and (IUGS), will require definition by GSSPs. Schindewolf Western Australia (but not New York) and of the entry of (1934) combined Divisions III and IV as the Prolobites- Manticoceras perhaps a little higher as shown in Western Platyclymenia Stufe. The Roman-numeraled divisions of Australia. UD I-D, the Sandbergeroceras Genozone, corre- Wedekind were retained in the classification developed by sponds to the first entry of the name genus and to the Mid- Becker (1993a, b), which is now subdivided using the dlesex Event. Roman alphabet, rather that the Greek as used by Wedekind UD I-E, the Probeloceras Genozone, corresponds to the and his followers (Becker & House, 2000). entry of the name genus and of Gogoceras and Uchtites. UD I-F, the Prochorites Genozone, corresponds to the entry of Cheiloceras Stufe, UD II, Nehdenian.–Now divided into that genus and probably a number of gephuroceratid genera nine divisions (UD II-A through I), of which the first six such as Carinoceras, Maternoceras, and Sphaeromanticoceras. (II-A through F) correspond to Wedekind’s II alpha, and re- UD I-G, the Mesobeloceras Genozone, corresponds to the mainder (UD II-G through I) corresponds to the former II entry first of Naplesites, then its descendant Mesobeloceras, beta, initiated by the entry of sporadoceratids. The Cheilo- and gephuroceratids such as Serramanticoceras and Costa- ceras Stufe equates with the Nehdener Schichten or Nehden- manticoceras. The base of UD I-H, the Beloceras Genozone, ian according to the original definitions of Paeckelmann corresponds to the point of derivation of the name genus (1924). from Mesobeloceras; in restricted environments Wellsites and The fauna of UD II-A, the Phoenixites Genozone, is an Schindewolfoceras are thought to belong here but improved impoverished one following the Upper Kellwasser extinc- correlation is desirable. UD I-I, the Playfordites Genozone, tions and is characterized by the opportunistic spread of corrsponds to the entry of the name genus and of Phoenixites, a genus first recorded early in the Frasnian (UD Timanoceras, Trimanticoceras, Virginoceras, and the pseudo- I-C). UD II-B, the Cheiloceras (Raymondiceras) Genozone, is ceratite Devonopronorites (dated by R. T. Becker on evidence characterized by the entry of simple-suture cheiloceratids from Iran). UD I-J, the Neomanticoceras Genozone, corre- with several species of Falcitornoceras. UD II-C, the Cheilo- sponds to the entry of the name genus and of Clauseniceras. ceras (Cheiloceras) Genozone, shows the entry of the true UD I-K, the Archoceras Genozone, corresponds to the spread Cheiloceras and the tornoceratids Polonoceras and Oxy- of the index genus with the Lower Kellwasser Event and Del- tornoceras. UD II-D, the Paratorleyoceras Genozone, shows phiceras; very early Crickites can occur here. UD I-L, the the entry of advanced cheiloceratids with pointed adventi- Crickites Genozone, corresponds to the acme of the name tious lobes (A), and includes Cheiloceras (Staffites), Parator- genus and the range of Crickites holzapfeli (Wedekind, leyoceras, Torleyoceras, and the tornoceratid Armatites. UD 1913), the zone fossil of Wedekind’s I delta; Enseites and II-E, the Praemeroceras Genozone, shows the entry of the Ceratobeloceras are limited to this level. All Gephurocerati- name genus. UD II-F, the Paratornoceras Genozone, shows dae, Acanthoclymeniidae, and Beloceratidae are extinct by the entry of oxyconic dimeroceratids such as Paratornoceras. the close of UD I-L, associated with the last stage of the UD II-G, the Maeneceras Genozone, shows the entry of spo- Upper Kellwasser Event. radoceratids, at first with only shallow A2 lobes. UD II-H, the Posttornoceras Genozone, marks the entry of the true Spo- Famennian radoceras with deep A2 lobes and of the name genus. UD II- Following Wedekind (1917, 1918), the Famennian has been I, the Dimeroceras Genozone, marks an international spread divided into ammonoid divisions II-VI, a practice con- of this genus (which appears rarely in UD II-H). firmed by Schindewolf (1923) and others. Originally these 78 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

Prolobites Stufe, UD III (with UD IV forming the Hem- ceras (Stockumites) but still with Cymaclymenia (Korn, 1991); bergian).–This is divided into three divisions. UD III-A, the this straddles the Devonian/Carboniferous boundary of the Pernoceras Genozone, corresponds to the entry of the new definition (Paproth et al., 1991). tornoceratids Pernoceras, Protornoceras, and Tornia. UD III- B, the Pseudoclymenia Genozone, is dominated by the GONIATITE BIOSTRATIGRAPHY Pseudoclymeniidae. UD III-C, the Prolobites Genozone, is OF THE LATE DEVONIAN IN NEW YORK characterized by the entry of many groups of clymeniids. The ranges of late Devonian goniatites in New York plotted against the stratigraphic divisions and the alignments of Platyclymenia Stufe, UD IV.–This is divided into three di- New York Regional Zones against the international goniatite visions, UD IV A-C, based on the entry of various clymenid zonation are summarized in Text-figs 23-24. This updates a genera. UD IV-A, the Prionoceras Genozone, is characterized previous review (House & Kirchgasser, 1993). For the Early by the Platyclymenia annulata (Münster, 1832) group, and and Middle Devonian, not covered in this report, there have marked by the entry of prionoceratids. UD IV-B, the Pro- been summary reviews in the past (House, 1962, 1978, toxyclymenia Genozone, is characterized by the name genus 1981b; Kirchgasser, 1985; Woodrow et al., 1989). So far as with early Cymaclymenia and Uraloclymenia. UD IV-C, the a contribution to international detailed zonation is con- Pachyclymenia Genozone, is marked by the entry of Protoxy- cerned, early work suffered from the problems associated clymenia (Franconiclymenia) and probably the earliest Platy- with an understanding of the complex facies relationships clymenia (Spinoclymenia), Pachyclymenia, and Costaclymenia. of the Devonian of New York State. For the Middle Devon- ian, the elucidation of the Hamilton sedimentary cycles and Clymenia Stufe, UD V, Dasbergian.–This is divided into their correlation in central and eastern New York by G. A. three divisions, UD V A-C, based on the entry of various Cooper (1930, 1933) was a special landmark. Although the clymenid genera. UD V-A, the Clymenia Genozone, is char- Tully Limestone had been long recognized as an important acterized by C. laevigata (Münster, 1832) and the spread of marker (Cooper & Williams, 1935), it disappears through Discoclymenia, Kosmoclymenia, Sphenoclymenia, Kiaclymenia, intra-Devonian erosion west of the Finger Lakes (Text-fig. Biloclymenia, and others. UD V-B, Ornatoclymenia Geno- 7). For the late Devonian, the classic work by Chadwick zone, is marked by the entry of O. ornata (Münster, 1834), (1935b) clarified the long-standing miscorrelations between with Gonioclymenia, Cycloclymenia, and distinctive species the eastern clastic facies and the more western neritic and of Kosmoclymenia. UD V-C, the Kalloclymenia Genozone, pelagic facies (Kirchgasser,1985). It might be true to say that is indicated by the entry of Piriclymenia piriformis (Schmidt, it was not until the fundamental review of Devonian corre- 1924), and of Kalloclymenia. lations by Cooper et al. (1942) that a stratigraphical frame- work comparable with European sections was available. By Wocklumeria Stufe, UD VI, Wocklumian.–This is divided then, Miller’s (1938) review of Devonian ammonoids had into six divisions, UD VI A-F, based on the entry of various been published, but that was a taxonomic, and not a bios- clymenid genera. UD VI-A, the Linguaclymenia Genozone, tratigraphic, treatment. is marked by the entry of Kosmoclymenia (Muessenbiaergia) Clarifications of the facies changes since 1950 have been bisulcata (Münster, 1840), Sphenoclymenia brevispina (Lang, fundamental to this work, especially the two correlation 1929), and Linguaclymenia, and corresponds to the Lower charts of L. V. Rickard (1964, 1975), which incorporated subarmata Zone and lower half of the Upper subarmata the results of correlations in the western New York Frasnian Zone of Korn (1986). UD VI-B, the Balvia Genozone, is at a very fine level. Also important has been the work of Sut- marked by the entry of the name genus and of various ton (1960, 1963), Sutton et al. (1962), Colton & de Witt species of Glatziella and was considered an upper part of the (1958), de Witt & Colton (1959, 1978), Pepper & de Witt Upper subarmata Zone by Luppold et al. (1994). UD VI-C, (1950, 1951), and Pepper, de Witt & Colton (1956). Of the Parawocklumeria Genozone, is marked by the entry of major significance to paleoenvironmental and sedimento- the name genus and of Kamptoclymenia and Triaclymenia; logical interpretation have been the many contributions by this is the Lower paradoxa Zone of Korn (1986). UD VI-D, G. Baird and C. Brett (e. g., Brett & Baird, 1996). the Wocklumeria Genozone, is indicated by the entry of An attempt at correlation of the New York sequence with Wocklumeria and of Synwocklumeria and Kielcensia. UD VI- Europe was made by House (1962). So far as an attempt at E, the Cymaclymenia Genozone, marks the remnant of cly- a local (New York) zonation is concerned, a start was made meniids after the extinctions of the Hangenberg Event. UD by House (1968) and continued in a review of the am- VI-F, the Acutimitoceras Genozone, is marked by the entry monoids of the Appalachians (House, 1978). In the latter of imitoceratids with evolute inner whorls, and Acutimito- work, numbers were given for faunas, and these are shown HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 79

Text-fig. 23. Ranges of late Devonian goniatites in New York State plotted against the stratigraphic divisions. Updated from House & Kirchgasser (1993). Near top, read Pharciceras? sp. 80 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

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). HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 81

in Text-figs 23-24. Since that time, international work by stone (House, 1978: faunas 7-12) and reviews have been the authors, usually jointly with R. Thomas Becker of the published (House, 1962, 1981b). New work by G. Kloc is University of Münster, Germany, has led to a much more not published; perhaps most significant is his discovery of detailed understanding of international faunas. As far as Maenioceras for the first time in New York, as the guide to New York is concerned, our work in Western Australia the Maenioceras Stufe and MD II. Agoniatites occurs and is (Becker et al., 1993) is particularly important. under study by S. Klovac (American Museum of Natural History, New York), and tornoceratids are abundant and REGIONAL NEW YORK GONIATITE ZONES were monographed by House (1965). The distinction of Zonation schemes are transitory in the sense that they are tornoceratids from below and above the Centerfield is im- subject to improvements and changes as new evidence be- portant because it seems that only late Hamilton tornocer- comes available. This section updates zonations published atids can be derived in the remanié and pyritic lags such as earlier (House, 1978; Kirchgasser & House, 1981; House the Leicester Pyrite. Huddle (1981) demonstrated the post- & Kirchgasser, 1993) and explains the correlation adopted Tully conodont ages of the pyrite lenses that also contain a with the international scheme of Becker & House (2000). derived goniatite fauna (Fauna 12) west of the last Tully out- This monograph covers the Upper Devonian in the classsical crops (Text-fig. 7). sense, that is, starting with the Tully Limestone. The new When the tornoceratid study was done, the Hungry GSSP redefinition of the base of the Frasnian Stage and Hollow Formation of Ontario was correlated with the Cen- Upper Devonian Series (Klapper et al., 1987; House, Feist terfield Limestone of New York, essentially following & Korn, 2000) has led to arguments on the taxonomy of Cooper et al., (1942). Just above the Centerfield, at Moon- conodont species thought to define the boundary (Johnson, shine Falls, Cayuga Lake, Polygnathus timorensis Klapper, 1989; Ziegler & Sandberg 1994, 1996; Sandberg et al., Philip & Jackson, 1970, is recorded, which indicates the 1988; Racki & Wrzolek, 1989; Racki & Bultynck 1993; conodont Lower varcus Zone (Klapper et al., 1970; Klapper, Klapper, 1988, 1989, 2000); some of this history has been 1981). Thus occurrences of Tornoceras uniangulare arkonense reviewed by House, Feist & Korn (2000). But, under the House, 1965, in the Ontario Arkona Shale (below the Hun- rules of the International Commission on Stratigraphy gry Hollow) and Tornoceras uniangulare widderi House, (ICS), the boundary is defined by the agreed point in the 1965 (Pl. 26, fig. 5) in the Ontario Widder Shale (above the GSSP at Puech de la Suque in southern France and not by Hungry Hollow) were thought to correlate to positions conodonts or goniatites. Some new regional zones are inte- below and above the Centerfield Limestone in New York. grated into the numerical system of House (1978) and Subsequently Brett & Baird (1985) questioned the Hungry Kirchgasser & House (1981) but further subdivided in the Hollow-Centerfield correlation, interpreting the Centerfield light of new evidence. Text-fig. 24 shows the New York Re- as a regressive unit within deeper water sediments and the gional Zones correlated with the conodont zonation scheme Hungry Hollow as the basal unit of a transgressive sequence. of Klapper (1989, 1997), following Becker & House (2000: Landing & Brett (1987) discovered a regional disconformity fig. 1, table 4); see also Klapper & Becker (1999) and House beneath the Hungry Hollow, but they interpreted the upper (2002). part of the Hungry Hollow and lower Widder Shale as an Following the methodology of House & Kirchgasser offlap-onlap cycle within the P. timorensis Zone and thus (1993) and Becker & House (2000), the Regional New York comparable to the upper part of the Centerfield. This issue Goniatite Zones are essentially defined by the first occur- was taken up by Sparling (1992) who found a fauna includ- rence of a single species or, in some cases, a group of species ing P.timorensis and P.ansatus Ziegler & Klapper, 1976, (the or fauna. guide to the Middle varcus Zone) in the Prout Limestone of Erie County, Ohio. Because the Prout has generally been Givetian considered of the same age as the Hungry Hollow, Sparling The Givetian Stage probably commences in New York a lit- used this presumption to argue that the Hungry Hollow is tle above the Cherry Valley Limestone and is indicated in also of Middle varcus Zone age. But the Prout fauna is rich, goniatite terms by the true tornoceratids (House, 1981b) and that of the Moonshine Falls locality is poor, and it could found in the Chittenango black shale of Cooper et al. (1942) be that the latter might still yield P. ansatus. More recently and Rickard (1975). This could be a local representation of Sparling (1999) has argued that, because the Tully has a late the Kacák Event (House, 1985, et seq.; 2002) . Middle varcus and younger conodont fauna, the Hungry Hollow should be correlated with the Tully Limestone. All Maenioceras Stufe, MD II (Faunas 7-12).–Several goniatite of this neglects the clear fact that the varcus Zone spans con- faunas have been found in the beds up to the Tully Lime- siderable time (House, 1995), and the Middle varcus Zone 82 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

could span from a level near the Centerfield to the early in the Lodi Limestone represents the latest Givetian (Middle Upper Tully (Text-fig. 7). As will be discussed later, the de- Devonian) in current definitions (Klapper et al., 1987), and rived goniatite faunas of the Leicester Pyrite are only of the the base of the Frasnian and Upper Devonian is taken where post-Hungry Hollow type. Thus Sparling’s correlation of the the Lower asymmetricus Zone (MN1) (or falsiovalis Zone) Widder Shale and Hungry Hollow with the Tully Limestone occurs at the junction of the Lodi Limestone and the over- is not accepted, although all could fall within part of the lying Penn Yan Shale (Kirchgasser, 1994). Middle varcus Zone (see Aboussalam & Becker, 2001: fig. 1; Aboussalam, 2003: fig. 2). Frasnian Manticoceras Stufe.–Because neither Neopharciceras or the Pharciceras Stufe.–This division was introduced by House critical species of Petteroceras are known in New York, it is (1985) for that part of the former Upper Devonian Zone of conodont evidence that places the Givetian/Frasnian bound- Pharciceras lunulicosta (G. & F. Sandberger, 1850) (I alpha ary precisely and it falls within our Fauna 15b. in Germany) that was transferred to the Middle Devonian Fauna 15c is marked by the entry of Chutoceras [C. with the redefinition of the base of the Upper Devonian nundaium, formerly Ponticeras cf. regale (Holzapfel, 1899)] (Klapper et al., 1987) by the International Commission on in the Ithaca Shale and Sandstone, at approximately the level Stratigraphy (ICS) and International Union of Geological of the Williams Brook Coquinite, approximately 53 m (175 Sciences (IUGS). Following work in Morocco, Becker & ft) above the Renwick Shale at Williams Brook (Loc. R, I- House (1994b) have established five zonal subdivisions 2) and 72 m (235 ft) above the base of the Renwick Shale (MD III A-E) as described earlier. in Fall Creek, Ithaca (Loc. X, Dy-10); ?C. nundaium occurs Regional Zone of Pharciceras amplexum (Fauna 13, MD at Cascadilla Creek (Loc. Z) and perhaps Enfield Glen (Loc. III-A).–The zonal species is only known from a single level BB). These Ithaca Formation occurrences of Chutoceras near in the Upper Tully Limestone (Moravia Bed or Bed G of the Cayuga Lake are equivalent in the west to the interval in the West Brook Member) as recounted earlier (see also House, Penn Yan Shale between the Lodi Limestone and the Crosby 1962, 1965; Kirchgasser & House, 1981) (Text-fig. 6). On Sandstone (Kirchgasser, 1985). These records are the most the eastern side of Seneca Lake, where the Tully is well de- precise evidence internationally of the entry of Chutoceras. veloped, P.amplexum occurs 0.38 m above the Bellona Coral Regional Zone of Koenenites styliophilus styliophilus Bed in Lodi Glen (Mill Creek, Seneca County; Loc. 3). At (Fauna 16a–b, UD I-B).–Koenenites enters in the upper June’s Quarry, Tully (Onondaga County; Loc. 7), the level Penn Yan Shale at the Linden Goniatite Horizon (Kirch- is 1.27 m above the Coral Bed. The genus occurs at several gasser & House, 1981) at Linden (Loc. 15) and is found more eastern localities as far as New Lisbon (Otsego eastward to Canandaigua Lake. It occurs with Tornoceras ar- County). See details under Tully Formation in the locality cuatum in the Crosby Sandstone (which is probably slightly section. younger than the Linden Horizon), near Keuka Lake. The Regional Zone of Pharciceras sp. (Fauna 14, MD III- entry of the genus defines the base of UD I-B. The genus is ?A).–The records of Pharciceras sp. from the Fillmore Glen last seen (as defined here) in the West River Shale. Two Bed of the uppermost Tully at Sheldrake and Mack creeks phases seem recognizable: and other localities (Loc. Gen-1) clearly postdate the P.am- Fauna 16a, the earliest phase, is dominated by Koenenites plexum horizon, but because none of the specimens known styliophilus styliophilus, which is restricted to the Upper Penn show sutures, they cannot be assigned. Yan Shale and is best known from the Linden Horizon and Regional Zone of Ponticeras perlatum (Fauna 15a-c, MD Crosby Sandstone and associated styliolinid beds. Acantho- III D-E/UD I-A).–There appear to be three local phases of clymenia (formerly identified as Probeloceras) enters in this this fauna. Fauna 15a is marked by the entry of Epitornoceras phase. (E. cf. mithracoides) in the upper Geneseo Shale, 12.2 m (40 Fauna 16b is characterized by Koenenites styliophilus kil- ft) below the Lodi Limestone, in Lodi Glen (Loc. 27), and foylei n. ssp. of the lower part of the Genundewa Limestone, approximately 24-27 m (80-90 ft) above the Tully Lime- especially at Bethany Center (Loc. 24/1) with Acanthocly- stone; the genus ranges to the middle Penn Yan Shale and menia genundewa and Tornoceras uniangulare compressum. equivalents. Koenenites? fasciculatus occurs at the type area of the Genun- Fauna 15b is marked by the entry of Ponticeras perlatum dewa Limestone at Canandaigua Lake (Loc. 23b). K.? fasci- in the upper Geneseo Shale, immediately below the Lodi culatus is also known from the Genundewa Limestone Limestone (House, 1962), for example in Hubbard’s Quarry equivalents near Keuka Lake but with conforming material (Loc. 27b); the Lodi Limestone yields the best preserved ma- from the earliest West River Shale (Locs DD, FF). terial (Kirchgasser, 1975). The conodont norrisi Zone fauna Regional Zone of Manticoceras contractum (Fauna 17a, HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 83

UDI-C).–Manticoceras enters in the upper Genundewa hypoxic transgressive phase. We have not searched the Mid- Limestone at Geneseo (Livingston County; Locs 19 and dlesex Shale systematically; it would appear likely that this 19a) and subsequently the genus is not seen until the upper genus is probably confined to a narrow band within the West River Shale. Together with M. contractum, two other Middlesex, perhaps at the time of maximum transgression. species were recorded from the Genundewa by Clarke We have found specimens of Sandbergeroceras in Middlesex (1898): M. nodifer and M. apprimatum; Acanthoclymenia cf. equivalents in Tioga County (Loc. 64). genundewa also occurs. This is the level that was formerly Regional Zone of Probeloceras lutheri (Fauna 19, UD I- taken to indicate the base of the M. nodulosum (Wedekind, E).–Typical faunas of the former Manticoceras cordatum 1913) (I beta) Zone (House, 1962). The entry of Mantico- Zone (I gamma) enter with the Cashaqua Shale in New York ceras represents a transgressive pulse. It is to be expected that although M. lamed aff. cordatum occurs earlier in the upper Timanites should precede this entry but there is no evidence West River Shale. In the lower Cashaqua Shale, P.lutheri en- of this form in the Appalachian Basin; that name is preferred ters and is common in the middle part and up to, but not for the division UD I-C, the Timanites Genozone, because including, the Shurtleff Septarian Horizon. M. sinuosum sin- it is a short-lived genus whereas Manticoceras continues until uosum occurs throughout the Cashaqua Shale and M. s. tar- the close of the Frasnian. In any case, the Genozone (or Di- dum and M. s. clausium n. ssp. are restricted to intervals in vision) terminology is one resulting from international, its lower part. Aulatornoceras eifliense is present in the middle rather than local, work. There is a gap in the record in the of the Cashaqua Shale. succeeding lower West River Shale. The type species for Regional Zone of Prochorites alveolatus (Fauna 20, UD I- Manticoceras, M. simulator (Hall, 1874), was recorded as F).–In the upper Cashaqua Shale, a new fauna appears that from the “Leiorhynchus Beds” of the Naples Formation at or is characterized by P.alveolatus, a senior synonym of P.strix near Ithaca, New York (see holotype and only known spec- Kirchgasser, 1968. Its entry is with the celebrated baritic imen, Pl. 9, Figs 4-5). The precise horizon in the Ithaca fauna of the Shurtleff Septarian Horizon, known best from Shale and Sandstone is not known but it is expected to be in Shurtleff’s Gully, Livonia (Livingston County; Loc. 41), the equivalents of the West River Shale, and probably high which yielded the type material of Acanthoclymenia neapoli- rather than low in that unit (Kirchgasser, 1985). tana (see House, 1961) but this form already appears slightly Regional Zone of Koenenites beckeri n. sp. (Fauna 17b, earlier. Manticoceras sinuosum sinuosum and Tornoceras uni- UD I-C).–K. beckeri n. sp., formerly referred to K. aff. lamel- angulare obesum are abundant in the Shurtleff Septarian losus (G. & F. Sandberger, 1851), occurs in the West River Horizon, and Aulatornoceras eifliense is also present. Shale, immediately below the Bluff Point Siltstone. One Regional Zone of Naplesites iynx (Fauna 21a, UD I- specimen is close to the specimen described from the Mon- G).–The horizon(s) of the type material for N. iynx (Clarke, tagne Noire under the name Hoeninghausia aff. archiaci 1898) and N. naplesense (Clarke, 1898) is not known but Gürich, 1896 (House et al., 1985: text-figs 8D-E, pl. 3, figs the specimens are said to have come from the Naples area of 10-12), but we decided that it is better to keep the genus Ontario and Yates counties. In view of the careful searching Hoeninghausia for undoubtedly oxyconic forms. The upper of the Cashaqua Shale, we feel that the horizon(s) must be West River Shale from the Bluff Point Siltstone to the top of in the lowermost Rhinestreet Shale. A major problem still re- the unit has horizons that yield koenenitids of variable qual- maining is the need for careful searching and correlation of ity of preservation and size, hence assignments are corre- levels yielding faunas in the Rhinestreet Shale, which is the spondingly difficult. Records include Acanthoclymenia aff. thickest black shale of the Frasnian black shales in New York neapolitana, Lobotornoceras aff. hassoni, and Manticoceras and thought to represent a considerable period of time. On lamed cf. cordatum. There is an evolute form that has been the international scale, it is known that Mesobeloceras (UD referred to as “Archoceras“ but that we now assign to Pon- I-G) is a later development of Naplesites but, as now inter- ticeras? sp. preted, that form is not known in New York and neither is Regional Zone of Sandbergeroceras syngonum (Fauna 18, the descendant Beloceras (which enters in UD I-H). MD I-D).–The zonal species is the only form known from Regional Zone of Wellsites tynani (Fauna 21b, UD- the Middlesex Shale and it has been widely reported from ?H).–This division approximately corresponds with the Be- New York and the Appalachians. S. syngonum was described loceras Genozone but we do not believe that true Beloceras is from Snyder’s Gully (Loc. 46c), Canandaigua Lake. This represented in New York. In its place is a curious group of multilobed and ribbed form is assigned to the Triainocerati- multilobed forms including Wellsites and Schindewolfoceras; dae and there is a long gap in the record of this group back these are found in the development of the Rhinestreet Shale to its presumed origin in the late Givetian. As with some in the central part of the state, especially south and southeast other multilobed genera, its occurrence seems related to a of Ithaca. Their occurrence well to the east and approaching 84 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

the more clastic facies of the Catskill Delta suggests that they (Becker & House, 2000). result from particularly extensive transgressions within the The occurrence of Playfordites cf. tripartitus in the upper major transgression represented by the Rhinestreet Shale and Rhinestreet Shale in the bed at Johnson Creek (Loc. 58/1) correspond to the triainoceratids in the Middlesex Shale in defines the New York Regional Playfordites cf. tripartitus representing unusual deeper-water forms. The facies is very Zone (22a), a zone that is restricted to that single level. The different from the Beloceras limestones of Europe and Aus- conodont fauna of the Reylea Creek Horizon, in the over- tralia. The Rhinestreet examples raise problems of their exact lying cycle, based on collections to be described with G. time equivalence with the western succession. Perhaps the Klapper, includes the marker species Palmatolepis semichato- earliest is the group represented by the genus Wellsites. W. vae Ovnatanova, 1976, which supports a correlation of the tynani is from near Elmira (Chemung County; Loc. 70), upper Rhinestreet P. cf. tripartitus occurrence with the con- and thought to be from the Moreland Shale, a basal tongue odont MN Zone 11. In the standard conodont zonation, of the Rhinestreet Shale. Wellsites williamsi is from Bald the level would be above the jamieae Zone, in the P.semicha- Mountain (Tioga County; Loc. 68), from a level probably tovae interval of the Lower rehenana Zone (Klapper & just above the Moreland Shale. Becker, 1999; Sandberg et al., 2002). Regional Zone of Schindewolfoceras chemungense (Fauna Playfordites cf. tripartitus (NYSM 12075) also occurs in 21c, UD I-?H-I).–This level marks the incoming of new the lower Angola Shale, in the Point Breeze Goniatite Bed forms with ribbed early whorls characteristic of the Triain- at Relyea Creek (Loc. 82/6), an anonomously high occur- oceratidae. The source of the original specimen of S. che- rence that we are placing, for reasons given below, in the mungense (Text-fig. 24I) was not precisely located but, when Neomanticoceras Genozone (UD I-J), that is, in the geno- it was refigured, Hall (1879: 69) gave the locality as zone above the Playfordites Genozone. Chemung near Owego (Tioga County; Loc. 69). Other Regional Zone of Sphaeromanticoceras rhynchostomum specimens referred to the genus with doubt are S.? equicosta- (Fauna 22b, UD I-J).–The goniatite faunas in the interval tum said to come from Athens, Pennsylvania, and later said from upper Rhinestreet Relyea Creek Horizon to the top of to come from a boulder there. The best localized specimen the Angola Shale define the New York Regional S. rhynchos- is S.? aff. equicostatum from Fairfield Forest (Loc. 67) near tomum Zone (22b), named for the most common mantico- Speedsville (Tioga County) at a level between the Moreland ceratid in the interval and particularly in the Relyea Creek Shale and Roricks Glen Shale tongues of the Rhinestreet Horizon of the upper Rhinestreet Shale and Point Breeze Shale, which Sutton & McGhee (1985) indicated to be still Goniatite Bed of the lower Angola shale. The placement of rather low in the Rhinestreet but above the levels bearing the S. rhynchostomum Zone in the Neomanticoceras Geno- Wellsites. zone or Division (UD I-J) and its differentiation from the Regional Zone of Playfordites cf. tripartitus (Fauna 22a, preceeding P. cf. tripartitus Zone (22a) is problematic: The UD I-I).–There is a thick interval, mostly of black shale, name genus for the Neomanticoceras Genozone has not yet within the Rhinestreet Shale that has yielded neither goni- been identified in New York, but already in the upper atites nor conodonts. However, in the upper Rhinestreet Rhinestreet Shale occurs the similarly oxyconic genus Shale in sections near Lake Erie, small-scale rhythms with Carinoceras, but that genus is internationally known to start calcareous levels or concretions occur and in this facies, and in UD I-F, that is, much earlier. A search is required of the in equivalents in Gardeau Shale and Sandstone farther east, New York oxyconic forms to see whether any develop the a suite of goniatite genera appear for the first time. At John- extra division in the ventral saddle that is characteristic of son Creek (Wyoming County; Loc. 58), in the concre- Neomanticoceras. tionary horizon (Loc. 58/1) in the cycle below the cycle with In the cycle above the cycle with the Playfordites horizon the Relyea Creek Horizon, a key marker species, Playfordites (Loc. 58/1) discussed above, is the most productive concre- cf. tripartitus (NYSM 12074) enters along with Carinoceras tionary level in the upper Rhinestreet Shale, the Relyea vagans, Manticoceras lamed, Sphaeromanticoceras rhynchosto- Creek Horizon, named for a bed at Relyea Creek (Loc. 60/3) mum, and Linguatornoceras aff. linguum. The genus Play- near South Warsaw (Wyoming County). This locality was fordites (type species: P.tripartitis) is the marker genus of UD known to Clarke (1899c) and possibly Luther (1903: 1012) I-I, the Playfordites Genozone or Division. Playfordites is not under the name Gibson’s Glen. The fauna of the Relyea known to occur outside of its genozone (or division) in Creek Horizon at Relyea Creek (Loc. 60/3) includes Western Australia, where the genozone was delineated Sphaeromanticoceras oxy, S. aff. oxy, Carinoceras vagans, Man- (Becker et al., 1993), or in Germany outside the classic ticoceras lamed, and, in the bed above (Loc. 60/4), S. rhyn- Manticoceras nodulosum Zone (I beta) of Wedekind (1913) chostomum and Linguatornoceras aff. linguum. Many of these or lower M. cordatum Zone of House & Ziegler (1977) forms continue upward into the lower Angola Shale where HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 85

Aulatornoceras auris, A. aff. eifliense, and Carinoceras sororium Lower Kellwasser Event (Feist, 1990) although Archoceras enter. wabashense (Kindle, 1901) is known from Indiana (House, In the sixth cycle above the base of the Angola Shale is a 1962) (Pl. 17, Fig. 1) along with Clauseniceras delphiense particularly rich level named the Point Breeze Goniaite Bed (Kindle, 1901) (Pl. 17, Figs 2-3). Detailed study has been (Kirchgasser & House, 1981) after a locality on Lake Erie undertaken of the early Hanover faunas by G. Kloc, and we Shore (Loc. 72/6). In addition to Playfordites cf. tripartites are indebted to him for some information, but his results from Loc. 82/6 discussed above, the Point Breeze Goniatite have not been published. The distinction between UD I-K Bed and the bed in the cycle below at Hampton Brook (Loc. and UD I-L in New York is not clear because the name gen- 74/6a) contain Sphaeromanticoceras oxy, Carinoceras vagans, era are not known [but see a record of ?Archoceras from Irish Manticoceras lamed, S. rhynchostomum, Linguatornoceras aff. Gulf (Loc. 90) referred to below]; the assignment to the linguum, A. aff. eifliense, and Tornoceras cf. typum (which be- genozones of Faunas 23 and 24 is therefore tentative. gins in Loc. 54/1 in the upper Rhinestreet), along with the Regional Zone of Delphiceras cataphractum (Fauna 23, first appearances of Aulatornoceras paucistriatum, Crasso- UD I-?K).–The first occurrence of the name species D. cat- tornoceras aff. crassum, and M. aff. lamed. The higher beds in aphractum in the lowermost Hanover Shale defines the base the Angola Shale are poor in goniatites. M. lamed and C. of the zone; it is a distinctive, small, rather evolute, and pe- aff. vagans occur in Loc. 79/10 at Varysburg and ?S. rhyn- riodically constricted manticoceratid that is not unlike Ar- chostomum occurs higher in the Angola above the “Trinity choceras varicosum (Drevermann, 1901) but has a subdivided Black” black shales at Loc. 18/14 near Varysburg (Gassman ventral lobe. In the absence of Archoceras at this level in New Road). York, a correlation with the Archoceras Genozone or Division The distinction between the Playfordites and Neomantic- (UD I-K) is problematic. oceras Genozones in New York is uncertain. Although Neo- Delphiceras cataphractum is common in the lowermost manticoceras has not been found in New York, we have Hanover Shale at Beaver Meadow Creek, Java (Wyoming placed the New York Regional Sphaeromanticoceras rhynchos- County; Loc. 92), where it occurs with Aulatornoceras tomum Zone (22b) of the upper Rhinestreet to the top of the rhysum in the first meter (Bed 92/1-2) of the Hanover Shale. Angola, in the Neomanticoceras Genozone or Division (UD At Walnut Creek, at Silver Creek (Chautauqua County; Loc. I-J). In doing so, we assume that the occurrence of P. cf. tri- 89a), the first records of D. cataphractum are approximately partitus in the Point Breeze Goniatite Bed of lower Angola 1.8 m above the Pipe Creek Shale and they continue over 10 shale represents an extension of the range of Playfordites m higher. The conodont age of the lowermost Hanover above its genozone into the succeeding Neomanticoceras Di- Shale is uncertain. It is assigned here to MN Zone 12 (Klap- vision (UD I-I). Comparison of the S. rhynchostomum Zone per et al., 1995) but to MN Zone 13 by Over (1997a, b), fauna with the upper Manticoceras cordatum Zone (I delta) corresponding to the Lower or Upper rhenana Zone in the fauna is not particularly helpful because most key marker standard zonation (Klapper & Becker, 1999). forms in Germany do not occur in New York, such as N. Regional Zone of Crickites lindneri (Fauna 24a, UD I- paradoxum (Matern, 1931a), Carinoceras galeatum L).–In the lower Hanover Shale at Walnut Creek (Loc. 89a) (Wedekind, 1918), Trimanticoceras cinctum (Glenister, C. lindneri enters the section near Bed 89a/4 or 5; the first 1958), Maternoceras sandbergeri (Wedekind. 1913) and rel- occurrence of this species defines the base of the New York atives, and Clauseniceras expectatum (Wedekind, 1913). Regional C. lindneri Zone and the International Crickites However, two forms in common, Crassotornoceras aff. cras- Genozone or Division (UD I-L). At Walnut Creek, other sum and Aulatornoceras paucistriatum, do occur in the Point loose specimens of C. lindneri probably came from Bed Breeze Goniatite Bed and these support an assignment of 89a/4 or 5. Bed 89a/5 is also the source of the specimen col- the New York Regional S. rhynchostomum Zone to the Neo- lected by G. Kloc that we earlier described as Sphaeroman- manticoceras Genozone (UD I-J) (Becker & House, 2000: ticoceras aff. rickardi (House & Kirchgasser, 1993: 247, figs. 129-130). The conodont fauna of the Point Breeze Goni- G-I). atite Bed is not diagnostic but it could already be in MN Regional Zone of Sphaeromanticoceras rickardi (Fauna Zone 12. The poor goniatites of the upper Angola are as- 24b, UD I-L).–The name form of this zone enters in the signed to the S. rhynchostomum Zone. lower part of the middle of the Hanover Shale (a loose spec- Regional Zone not assigned (Fauna 23, UD I-K).–No imen from near Bed 91/1 at Glade Creek, north of Stryk- fauna is known from the Pipe Creek Shale that we consider ersville, Wyoming County) and ranges to the upper to represent the Lower Kellwasserkalk level of Europe. Also Hanover where a single specimen was found by L. Rickard in New York, there are no records of Archoceras, a genus that in Bed 91/7 in Glade Creek, 7.6 m (25 ft) below the base of in Europe becomes opportunistically abundant with the the Dunkirk Shale (Text-fig. 16). New York Faunas 24a and 86 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

24b together correlate with the lower subzone of the Crick- Famennian ites Genozone as defined by Becker & House (2000: 132). Regional Zone Unnamed to top of Frasnian Stage Cheiloceras Stufe.– (Fauna 24c, UD I-L).–Near the top of the Hanover Shale in Regional Zone of Cheiloceras (Cheiloceras) amblylobum Bed 90/2 in the section at Irish Gulf, southeast of North (Fauna 25, UD II-C).–The most significant level with this Boston (Erie County), small goniatites with distinctively fauna is the Corell’s Point Goniatite Bed of the Gowanda evolute and compressed inner whorls determined as ?Ar- Shale, traceable from Lake Erie Shore at Corell’s Point (Loc. choceras sp. (NYSM 16587 and 16588) occur with small 97), and generally eastward at several localities as far as subinvolute goniatites with a rounded, rather rectilinear Marshfield (Erie County; Loc. 106). Cheiloceratids referred whorl section assigned as ?Crickites sp. juv. (Text-fig. 45H; to the name species continue eastward at least as far as Java NYSM 16589). The whorl section of ?C. sp. juv. is similar (Wyoming County; Loc. 109) and probably farther to the to that of C. holzapfeli, the highest Frasnian goniatite zone- east. Fauna 25 comprises C. (C.) amblylobum, Truyolsoceras fossil in Europe. The goniatites of Bed 90/2, constituting bicostatum, and Phoenixites concentricus. The name genus Fauna 24c, are the highest known goniatites in the New confirms correlation with the Cheiloceras (C.) Genozone UD York Frasnian and they define an unnamed Regional Zone. II-C (Becker & House, 2000). Fauna 24c might correlate with the upper subzone of the Regional Zone of Truyolsoceras clarkei [Fauna 26, UD II- Crickites Genozone as defined by Becker & House (2000: ?C (?II-B)].–There is an early level in the Gowanda Shale 132). with T. cf. bicostatum in Walnut Creek (Loc. 99a) near Bed 90/2 is a 2.0-2.5 cm-thick baritic, crinoid and gas- Forestville (Chautauqua County); the same village is men- tropod-rich, concretionary horizon that is 4.7 m below the tioned for the locality of the types of the very different form base of the Dunkirk Shale and 1.76 m below the conodont- described here as T. clarkei. Note that earlier (House 1962, defined Frasnian/Famennian boundary (Over, 1997a, b). It 1965), Miller’s species was referred to Aulatornoceras. The is shown in Text-fig. 16 (2 cm bed) and by a concretion sym- exact level of the T. clarkei deposit has not been determined bol in Over’s (1997b) fig. 8. The conodont fauna of Bed but it is not thought to be so high as to be described as near 90/2 indicates the highest Frasnian MN Conodont Zone 13 the Laona Siltstone. Further collecting is needed at these lev- (Over, 1997b: fig. 8) and conodont linguiformis Zone of the els. The queried reference to Genozone UD II-B has to be standard zonation (Klapper & Becker, 1999). quite tentative in view of the absence of simple-sutured No higher Frasnian goniatites are known in New York, cheiloceratids. but possible goniatite anaptychi (ammonoid opercula) occur Regional Zone of Maeneceras aff. acutolaterale (Fauna 27, in the Huddle Rider Bed, a thin black shale a meter or so UD II-G).–No representative of Oxytornoceras, Praemero- below the top of the Hanover in Glade Creek (Loc. 91, Text- ceras, or Acrimeroceras, all guides to UD II-D-F, are known fig. 16) and approximately 2 m below the Frasnian-Famenn- in New York or North America. The next level, the ian boundary. The conodont-defined Frasnian-Famennian Maeneceras Genozone UD II-G, is represented by a single boundary, as determined by Over (1997a, b) and shown in specimen (USNM 137645) of M. aff. acutolaterale, previ- Text-fig. 16, lies within the upper Hanover Shale or lower ously determined as M. cf. pompeckji, from the middle part Dunkirk Shale (Locs 91 and 92) depending on where the of the Ellicott (Chadakoin) Shale near Summerdale (Chau- Hanover-Dunkirk boundary is drawn. tauqua County; Loc. 110). R. T. Becker informed us that Homoctenids have been noted near the Hanover- there are problems regarding the type specimen of M. pom- Dunkirk boundary at some localities and these distinctive peckji, in which case that name would be replaced by M. dacryoconarids were once thought to be extremely rare after acutolaterale. These forms with shallow A2 lobes have been the end-Frasnian [Schindler (1990) recorded only a single investigated by Bochwinkle et al. (n. d.), who referred the specimen above the Upper Kellwasserkalk], but now they are species to UD II-I, but the New York form is likely to be known to range well into the Famennian, for example, in earlier. China (Xing Li, 2000). At Beaver Meadow Creek (Loc. 92), Fauna 27 is the highest known in New York, but it is homoctenids occur profusely in the first siltstone above the convenient to carry the known succession of eastern North Frasnian-Famennian boundary (Over, 1997a, b; Over et al., America to the top of the Devonian by commenting on the 1997); this horizon is the first of the three stippled units in faunas of Pennsylvania and Ohio. the lower Dunkirk Shale shown by Pepper & de Witt (1951: Regional Zone of Maeneceras milleri (Fauna 28, UD II- Section 7, Java). Homoctenids are also abundant in black H).–The name form, a single specimen, has been described shale 50 cm above the Frasnian-Famennian boundary in by Flower & Caster (1935), Miller & Flower (1936), and Glade Creek (Loc. 91). Miller (1938). If Sporadoceras is defined by its pointed A2 HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 87

lobes, rather than the depth of the A2 lobes, as proposed by haps UD VI-F. Bockwinkel et al. (1999), then Maeneceras is the correct as- UMMARY signment. But the size at which the pointed A2 lobe is S achieved is essential to the definition. The specimen is from Thus there is represented in New York and adjacent states an Howard Quarries (Erie County; Loc. 111; White, 1881: excellent succession of late Devonian ammonoid faunas. Be- 103), Pennsylvania, beside Falls Run, approximately 8 mi cause of the thick nature of the succession, stratigraphical (12.8 km) from the Lake Erie Shore. The Lower Venango precision is as fine as anywhere in the world. Only a few lev- (Cattaraugus) Sandstone crops out just below the quarry. els are rich, however, and those are mostly in the Frasnian. The Regional Zone of M. milleri correlates to the Post- In general, the faunas are of low diversity and abundance tornoceras Genozone UD II-H of Becker & House (2000). and this is thought to be because of the restricted environ- ment provided by the Appalachian embayment in which Prolobites and Platyclymenia Stufe.– they occurred, an environment that probably did not have Regional Zone of “Pleuroclymenia” ohioensis (Fauna 29, regular access to pelagic environments in which most De- UD IV-A).–No faunas equivalent to the Prolobites Stufe are vonian ammonoids flourished, as was long ago noted by known, unless ?Genus edwinhalli (Clarke, 1898), a species Clarke (1902). The individual horizons at which faunas that in this report is unassigned, is included. The type local- occur, however, might well represent the occasions when ity for the P.ohioensis Zone level is on the West Branch of the small rises of sea level enabled ammonoid faunas to enter Vermilion River, 550 yd (490 m) northwest of the NY Rte. the basin, and in the future these marine bands might serve, 113 road bridge over the river at Birmingham, 12 mi (20 as in the Carboniferous, as ecological as well as chronostrati- km) west of Cleveland, Ohio. The fauna comes from the graphic markers. upper part of the lowest 1.8 m of the Cleveland Shale below Some discussion of the transgressive pulses that seem rec- a siltstone unit and includes Platyclymenia spp., P. ohioensis ognizable in the New York succession and to which the entry House, Gordon & Hlavin, 1986, Cyrtoclymenia, and of new goniatite faunas, or marker bands, seem related has Maeneceras inflexum (Wedekind, 1908) (House et al., 1986). been documented elsewhere (House 1983; House & Kirch- Fauna 29 appears to be early in the Platyclymenia Stufe, or gasser 1993; House et al., 2000) and has been also covered, Prionoceras Genozone UD IV-A (Becker & House, 2000). especially with relevance to dating of events by conodonts, by Johnson et al. (1985, 1986), Klapper et al. (1995), and Clymenia and Wocklumeria Stufe.– Klapper & Becker (1999). Regional Zone of Cymaclymenia sp. (Fauna 30, UD VI- The faunas reported here are from a considerable area E).–No goniatite faunas equivalent to the Clymenia Stufe and from a substantial thickness of strata. Clearly, we have (UD V) are known in eastern North America. There is a not exhausted knowledge of the ammonoids of this classic level near Rockford, Indiana, which has yielded a specimen and beautiful area, but we hope the time framework laid referred to Epiwocklumeria? sp. that might correspond to the down in this work will enable others to fill in the many miss- Wocklumeria Genozone UD VI-D (Becker & House, 2000) ing gaps that doubtless remain. but that cannot yet be placed in the Ohio succession (House et al., 1986). SYSTEMATIC PALEONTOLOGY From 0.6-1.5 m (2-5 ft) below the top of the Cleveland Shale on Chance Creek (Lorain County), Ohio, and some INTRODUCTION other localities, Cymaclymenia sp. occurs as flattened pyritic In the taxonomic descriptions that follow, lengthy syn- films together with poor indeterminable sporadoceratids and onomies are not included and concentration is made on the prionoceratids (House et al., 1986). Fauna 30 correlates to description of type material. Within the families, the nom- the Wocklumeria Stufe, Cymaclymenia Genozone UD VI-E inate genus is described first and other genera are treated in (Becker & House, 2000). approximate stratigraphical order. The same pattern is fol- Regional Zone of Rectimitoceras quadripartitum (Fauna lowed within genera as far as possible. Abbreviations of mor- 31, UD VI-?F).–Faunas near the base of the Bedford Shale, phological terms used are as given in Text-fig. 25 or are for example, near Bedford (Cuyahoga County), Ohio, 1.7 explained as they are used. The general terminology for shell mi (2.7 km) northeast of Amherst City Hall, and Tinkers form, sutures, and growth lines has changed little since the Creek and neighboring localities (House et al., 1986), yield accounts of Miller (1938) and Moore (1957), but more de- a fauna including R. (Prionoceras) quadripartitum (Münster, tailed accounts of sutural terminology are available (Schin- 1939) without any other definitive Famennian forms. The dewolf, 1954; Kullmann & Wiedmann, 1970; Wiedmann fauna is thought to be high in the Wocklumeria Stufe, per- & Kullmann, 1981; House, 1996). 88 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

The section Distribution gives the stratigraphic and ge- the NYSM or the Paleontological Research Institution ographic distribution of each taxon. Under the heading (PRI), Ithaca, where the Cornell collection retains its origi- Range, the biostratigraphic position of each taxon is listed in nal numbering. The PRI also holds much other material. the following order: (1) name of Stage (e. g., Givetian), (2) Some early material of H. S. Williams and material collected genus name and key of International Genozone or Division by G. A. Cooper is in the U. S. National Museum (USNM), [e. g., Pharciceras Genozone MD (Middle Devonian) III-A] Washington, DC (now National Museum of Natural His- following Becker & House (2000), (3) name of New York tory), with some material collected more recently. The avail- Regional Zone and Fauna number in Appalachian Basin able material collected by E. M. Kindle is held in the succession [e. g., Pharciceras amplexum (13)] following Geological Survey of Canada, Ottawa (GSC). Some speci- Kirchgasser & House (1981), House & Kirchgasser (1993), mens referred to are in the Buffalo Museum of Science and Becker & House (2000), and (4) name of Standard (BMS), the geological collections at the University of Michi- Conodont Zone (e. g., Middle varcus Zone) following Sand- gan, Ann Arbor (AAM), the Deparment of Geoscience at berg et al. (1989), Ziegler & Sandberg (1990) and Klapper the University of Iowa, Iowa City (SUI), the Sedgwick Mu- & Johnson (1990), and, for the Frasnian Stage, Zone Num- seum, Cambridge (SM), the University Museum, Oxford ber of Montagne Noire (MN) Conodont Zone following (OUM), the University of Western Australia, Perth (UWA), Klapper (1989, 1997), Klapper et al. (1995), and Klapper & the Musée royale d’Histoire Naturelle, Brussels (IG), the Becker (1999). Museum für Naturkunde der Humboldt Universität, Berlin (HUM), and the Museum für Naturkunde, Berlin (HM REPOSITORIES MBC). The major collections referred to, including the bulk of Hall The material resulting from our earlier work (House, and Clarke’s material from the nineteenth century, are in the 1962, 1965; Kirchghasser, 1975) has already been deposited New York State Museum, Albany (NYSM), which is where in the NYSM and cataloged (Kilfoyle, 1969). The bulk of most of our own collections have been deposited. Some early the material collected after 1965 is deposited in the NYSM; material is in the American Museum of Natural History in this material was initially labeled in green paint with black New York (AMNH). The large collection, mainly by H. S. locality numbers in the 3000s and each specimen in an in- Williams and especially J. W. Wells, formerly at Cornell dividual collection numbered (e. g., 3136/9). A catalog of University Museum of Paleontology (CU), is now either in these numbers has been deposited with the NYSM. Speci- mens figured, or referred to, have been assigned NYSM numbers. Other fossil material is deposited in the NYSM stratigraphical collection. Copies of our field notes, section logs and catalogs are also deposited in the NYSM.

Phylum MOLLUSCA Cuvier, 1897 Class CEPHALOPODA Leach, 1817 Subclass AMMONOIDEA Agassiz, 1847 Order ANARCESTIDA Miller & Furnish, 1954 Superfamily PHARCICERATOIDEA Hyatt, 1900 This superfamily is the first of the two major stocks showing a progressive proliferation of umbilical lobes on the flanks; the second is the Gephuroceratoidea. Both characterize, or rather, define, the German Adorfian as originally used (Wedekind, 1913). The families included in the Pharcicer- atoidea are the Pharciceratidae, Petteroceratidae, Eobelocer- atidae, Devonopronoritidae, and Triainoceratidae. Only the first and last are represented in New York State. The Pharciceratidae and Triainoceratidae are well-de- fined groups that are discussed in further detail below. Both show a morphological series of forms of successive sutural complexity, the distinction between them resting on the co- station and nodation of the latter. The Eobeloceratidae is a Text-fig. 25. Diagrams illustrating terminology used in goniatite late Givetian group that does not show the typical pinched descriptions and measurements given in the text. HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 89

lateral lobes of the Pharciceratidae. The Givetian Petterocer- tilobed forms, which could comprise these, from those with atidae, which just reaches the lowest Frasnian, differs from fewer lobes like Pharciceras. Too little is known, however, to all other families in the initiation of additional lobes at the follow such a course at present and the Synpharciceratidae venter. The Devonopronoritidae (Bogoslovsky, 1958; 1969: is regarded as a junior synonym of Pharciceratidae. 290-295) contains solely Devonopronorites, an endemic odd- Distribution.–Representatives known in Europe, North ity from the Rudnyi Altai, Russia, with a bifid first lateral Africa, Russia, China, and eastern North America (Becker & lobe, an interesting approach to ceratitic frilling. House, 2000). Here the name Pharciceratoidea is used in preference to Range.–In Europe, the family is restricted to late Givet- Triainocerataceae essentially following A. K. Miller & W. ian to earliest Frasnian: Pharciceras to Neopharciceras Geno- M. Furnish (in Moore, 1957: L33), although the recogni- zones MD III-A-E to UD I-A. Formerly taken to be tion of the suborder Gephuroceratina rearranges the status. restricted to the long established lunulicosta Zone (lower- The superfamily Triainoceratoidea has been used by Bo- most Manticoceras Stufe), or Oberdevon I alpha, which is a goslovsky (1969: 113) essentially as a replacement name for division now mostly in the Middle Devonian following the Pharciceratoidea, which he had also used on earlier occasions decision of the Subcommission on Devonian Stratigraphy (in Orlov, 1962: 341). The status of Triainocerae (Hyatt, to raise the Middle/Upper Devonian boundary to a level 1884: 333) is rather questionable for taxonomic purposes, above the range of the family (Klapper et al., 1987). and the form in which the name was introduced (within brackets rather than without as used for the clear designa- Genus PHARCICERAS Hyatt, 1884 tions) implies that it was not envisaged as formal higher- Type species.–Goniatites tridens G. & F.Sandberger, 1850, taxon characterization (indeed, some groups proposed thus by subsequent designation (Wedekind, 1918: 127). comprise generic names only). It seems reasonable to prefer Diagnosis.–Pharciceratids with varied shell form, from the unquestionably correct derivation of Pharciceratoidea. widely to narrowly umbilicate and well-rounded whorl sec- Notice, however, that the same comment could be made for tion at least in early whorls. Growth lines biconvex with lap- Triainoceratidae, which is more recent, in its correct form, pets often associated with ventrolateral furrows. Suture with than Sandbergeroceratidae (Miller, 1938: 24, nom. transl. a simple mid-ventral lobe, a first lateral lobe, in many Ruzhencev, 1957: 55), so that there is a certain illogicality smaller than the second lateral lobe, and one to three lobes in the procedure adopted here, the intention of which is to between the second lateral and the umbilical seam that form disturb terminology as little as possible. as umbilical lobes during ontogeny. In New York, only the late Givetian Pharciceratidae and Included species.–Pharciceras tridens (Sandberger & Sand- the Frasnian Triainoceratidae are known. The Eobelocerati- berger, 1850), P.amplexum (Hall, 1886), P.bidentatum Pet- dae, Petteroceratidae, and Devonopronoritidae are not ter, 1959, and P. galeatum Wedekind, 1918. known to be represented in North America. Discussion.–With the creation of Stenopharciceras by Montesinos & Henn (1986), the genus is now restricted to Family PHARCICERATIDAE Hyatt, 1900 forms with rather few lateral lobes. The simplest forms, Diagnosis.–Pharciceratoideans with 3 to at least 23 lateral Pharciceras tridens and P.amplexum, seem to be earliest; the lobes (including those on the seam) in the adult with simple former is the earliest recorded in North Africa (Bensaïd, to trifid ventral lobes. Growth lines biconvex. Form sub- 1974) and the latter, in New York, has the earliest conodont globular to subevolute, outline usually rounded, rarely oxy- date (late Middle varcus Zone). conic, mostly without ribs and nodes. Distribution.–Pharciceras is known from eastern North Discussion.–This family includes a wide range of distinct America (Tully Fomation and equivalents between Seneca types resulting from a radiation during the late Givetian fol- and Otsego counties, New York), Europe (Spain and Ger- lowing initiation of forms with additional umbilical lobes, many), and North Africa (see Addendum). the characteristic of distinction from the supposed ancestral Range.–Givetian: Restricted to Pharciceras Genozone Maenioceratidae (House, 1978: 45). MD III-A, which is defined by its appearance. Appears to The family includes the genera Pharciceras Hyatt, 1884 range from top of conodont Middle varcus Zone to disparilis (= Sphaeropharciceras Bogoslovsky, 1955b: 1104), Neophar- Zone but range is uncertain, as conodonts are rare or absent ciceras Bogoslovsky, 1955a: 95, Stenopharciceras Montesinos at many of the localities. & Henn, 1986, and Synpharciceras Schindewolf, 1940: 430. It has been reviewed by Bogoslovsky (1969). Schindewolf Pharciceras amplexum (Hall, 1886) (1940) distinguished a family, the Synpharciceratidae, and Pl. 1, Figs 1-8; Text-figs 26D-H; Table 2 this might prove useful if it is ever needed to separate mul- 90 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

Table 2. Biometric data for Pharciceras amplexum (Hall, 1886) from the Tully Limestone. Here, and in all tables of biometric data to follow, D = diameter, U = umbilicus, UW = umbilical width, Wh = internal whorl (aperture) height, WH = whorl height, WW = whorl width (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 0.57 USNM 96545a 39.6 12.3 12.5 18.0 0.98 0.45 USNM 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 0.43 USNM 143014 ca. 30.5 11.0 10.0 ca. 13.3 1.10 0.44 SUI 37052c ca. 29.0 ca. 11.7 — 12.8 — 0.44 USNM 137661 29.0 9.7 10.0 12.3 0.97 0.42 NYSM 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 0.43 SUI 37052c 13.1 7.5 ————

Goniatites amplexus Hall, 1886, pl. 127 (12), fig. 1; Beecher in follow course of growth lines, seen on flanks only, number- Hall & Clarke, 1888: 39-40, pl. 127, fig. 1. ing 24 in previous half whorl at 13.6 mm diameter (SUI “Goniatites” amplexus. Cooper & Williams, 1935: 859. 37052b) but coarser in other specimens (NYSM 12001). Maeneceras amplexum. Clarke & Ruedemann, 1903: 602. Suture as illustrated (Text-figs 26D-F); dorsal suture un- Frech, 1913: 23. Manticoceras amplexum. known. Manticoceras? amplexum. Miller, 1938: 74-76, pl. 13, figs 15-19. Pharciceras amplexum. House, 1962: 256, 272-274, text-figs 10C- Discussion.–Substantially more material is now available D, pl. 45, figs 1-4, 7-9; 1968: 1065; Bogoslovsky, 1969: 285; on which to describe the species. The material is not well House & Ziegler, 1977: 90, text-fig. 91; House, 1979: 54, preserved and although commonly showing sutures and text-figs 9G-I; House & Kirchgasser, 1993: 276. growth lines, detailed elucidation of ontogeny has not been possible. Many specimens are distorted as result of fractur- Type material.–The holotype (NYSM 3729), the only ing. Hence some of the measurements are approximate. This specimen known to Hall when he illustrated the species, was species differs from openly umbilicate maenioceratids only described by Beecher (1888) and by Miller (1938) and is in the additional lobe near the umbilical seam. It is the ad- reillustrated here (Text-fig. 26F, Pl. 1, Fig. 3); it is a small dition of one or more extra lobes in this position that is the septate specimen showing growth lines. generic distinction of Pharciceras from Maenioceras. The Additional material.–In addition to the material de- species is very close to the lectotype, here designated, of scribed by House (1962, USNM 143014, 137661, 96545a Pharciceras tridens (G. & F. Sandberger, 1850: pl. 4, fig. 2; and b, 96552), material listed below now in the NYSM and 1851: 66; and refigured here, Pl. 1, Fig. 10). As can be seen, SUI 37052a,b,c. however, from the dimensions of the lectotype of P. tridens Dimensions.–See Table 2. previously published (House, 1962: 273), that species is Description.–Shell form subevolute; whorl outline in more depressed at comparable diameters, although SUI early whorls depressed with well-rounded flanks, weak ven- 37052a approaches it in certain respects. German material trolateral spirals, and tabular venter. At approximately 25 of P. tridens (e. g., see Kullmann & Ziegler, 1970: text-fig. mm diameter, whorl height equals whorl width and approx- 2A, pl. 1, fig. 3) shows that the type species can be consid- imately equidimensional outline continues to largest diam- erably larger and in another specimen, illustrated by the eters seen (USNM 96545a), but WW/WH ratio varies. In Sandberger brothers (1851: pl. 9, figs 2, 2a, 2b; refigured larger specimens, ventrolateral groove better developed, by House & Ziegler, 1977: pl. 5, figs 24-26), the sutural el- often double (USNM 96545b; NYSM 11996 at 31 mm di- ements seem more advanced at comparable diameters. Both ameter) and well marked; less so in other cases (SUI characters might suggest a later derivative and younger 37052c). Growth lines biconvex, passing slightly forward species than P. amplexum. over umbilical wall, then backward to broad, rounded sinus The entry of Pharciceras was taken in Germany to mark on mid-flanks, then sweeping forward into ventrolateral fur- the base of the Adorfian and Upper Devonian (Frech, row to form lappet and back into deep ventral sinus; lappet 1887). The attempt of Kullmann & Ziegler (1970), how- relatively longer in earlier stages. Very fine ribbing developed ever, to avoid assignment of P. amplexum to Pharciceras has on flanks before approximately 15 mm diameter: fine ribs been precluded by the evidence from excellent material in HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 91

Text-fig. 26. Sutures and cross sections of Pharciceras tridens (G. & F. Sandberger, 1850) from Germany and the sutures and cross section of P.amplexum (Hall, 1886) from the Tully Limestone of New York. A-C. Pharciceras tridens. A. SM H 9935, suture of a specimen from the Untere 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 have been developed by abrasion that has resulted in the rounding of some of the lobes. D-H. Pharciceras amplexum. D. SUI 37052, suture of a specimen from June's Quarry, Tully (Loc. 7) drawn by W. M. Furnish, X 3. E, USNM 143014, suture based on a specimen from the same locality, X 3. F. NYSM 3729, suture based on the holotype from near Lodi Landing (Loc. 3), Seneca Lake, Seneca County, X 3. G, NYSM 12002, 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.

the Iowa collection (House, 1978: 554). House (1962: 273) figs 5-6) of the Tully Limestone (Heckel, 1973) or the upper drew attention to the affinities of P.amplexum to Triainoceras part of Bed G in the West Brook Member of Cooper & but the lack of marked ribbing precludes reference to the Williams (1935: 791). From Lodi Glen, Seneca Lake Triainoceratidae. Kullmann & Ziegler (1970) also consid- (Seneca County; Loc. 3), the most westerly known record, ered Tamarites (Bogoslovsky, 1969: 275), an evolute ribbed comes the holotype (NYSM 3729) and other specimens form that has a similar subdivision of the ventral lobe (but (NYSM 12002a and b, 12003) one of which was collected simpler overall suture), and used evidence from G. A. in situ by J. W. Wells. From June’s Quarry, Tully (Onondaga Cooper’s comments on brachiopods associated with County; Loc. 7), comes USNM 143014, SUI 37052a-c, Tamarites to argue that such types could be Middle Devon- and NYSM 11996-11999, 12004, 12005). From Tinkers ian – all without drawing attention to the fact that in Falls (Onondaga County; Loc. 8), comes USNM 137661. Cooper’s view of the placing of the Middle/Upper Devonian Also from the West Brook Member, but at localities far- boundary (Cooper et al., 1942; Cooper, 1968), any I alpha ther east, are several specimens referred here that were col- goniatite would be assigned to the Middle Devonian because lected during the survey by Cooper & Williams (1935). The he took the Genundewa Limestone as the base of the Upper precise correlation of these with the Pharciceras-bearing level Devonian. All this has been resolved by the decision of the farther west can only be inferred. From near Lebanon International Union of Geological Sciences (IUGS) and (Madison County; Loc. 9c), is a single specimen (USNM Subcommission on Devonian Stratigraphy (SDS) to define 96552). From near Laurens (Otsego County; Loc. 9b), are the Middle/Upper Devonian boundary at a much higher several specimens (USNM 96545a and b). This locality near level than the Tully Limestone (Klapper et al., 1987). Laurens is the most easternly record of P.amplexum known. Distribution.–Moravia Bed or West Brook Member of There is no other record of this species elsewhere in eastern the Tully Formation between Seneca and Cortland counties. North America of which we are aware. Molds of probable Almost all of the known specimens of Pharciceras amplexum Pharciceras occur in some abundance in the Filmore Glen certainly come from a single level in the Moravia Bed (Text- Bed of the Tully and rarely in the overlying Geneseo Shale 92 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

(Pl. 31, Figs 14-15). boniferous Pseudarietites is unrelated to these other genera Range.–Givetian: Pharciceras Genozone MD III-A. Re- and is derived from Gattendorfia in the early Tournaisian gional Zone of P.amplexum (13). Conodont late Middle to (Weyer, 1972). Miller (1938: 179) thought that Sandberge- Upper varcus Zone (Klapper, 1981: fig. 2). See also Abous- roceras sandbergerorum Miller, 1938, from Oberscheld and salam & Becker (2001: fig. 1) and Aboussalam (2003: fig. Königzug, Germany, shows evidence of an incipient trifid 2). lobe, but now that the type has been photographically illus- trated for the first time (House & Ziegler, 1977, pl. 6, figs Family TRIAINOCERATIDAE Hyatt, 1884 16-18) there is really no strong evidence for this. Certainly Diagnosis.–Pharciceratoideans with four to perhaps ten there is nothing to approach the marked trifid lobes of T. or so ventral lobes in adult with ventral lobe small and sim- gerassimovi and T. costatum (d'Archiac & de Verneuil, 0000), ple to large and trifid. Shell form with strongly costate, evo- so that the generic distinction is worth keeping. lute inner whorls, smoother outer whorls, and tendency The material described here from New York is mostly toward compressed or oxyconic form in adult. poorly preserved, very little of it showing evidence of su- Discussion.–The species included in the Triainoceratidae tures. It appears, however, that shell ornament gives some have markedly ribbed inner whorls but costation is virtually consistent stratigraphical information. None of the material lost in adults. Almost all species show ventrolateral furrows assigned to Sandbergeroceras shows a complete suture or ev- at some stage, and they are often double. There is a charac- idence of the ventral lobe. Forms assigned to Sandbergero- teristic increase in the size of lobes from the umbilical seam ceras occur early, apparently in the Sonyea Group, probably toward the venter. The critical distinction between the in- beginning in the Middlesex Shale. They are characterized by cluded genera, as presently understood, lies in the form of ribbing in the inner whorls that is not known to extend be- the ventral lobe. Much has to be learned concerning the yond 30-40 mm diameter, with individual ribs prorsiradiate family and, apart from Triainoceras gerassimovi Bogoslovsky, to approximately rectiradiate (as in the type species of the 1958, none of the included species can be said to be well genus, S. sandbergerorum); fine growth-line lirae often lie be- known. tween the ribs. Specimens referred to Schindewolfoceras, on The family Triainoceratidae (nom. correct., Ruzhentzev, the other hand, might begin in the upper Sonyea Group, 1960: 175, pro Triainocerae Hyatt, 1884: 336) comprises but are characteristic of the West Falls Group. In the latter, the four genera Triainoceras, Sandbergeroceras, Wellsites, and the ribbing usually extends to at least 80 mm diameter, and Schindewolfoceras; all genera are confined to Oberdevon I of the course of the ribs is rursiradiate over the outer flanks and the traditional German definition. To these four genera is often also distinctly backwardly directed over the lower tentatively added Tamarites Bogoslovsky, 1965, the strongly flanks. This is no more than an initial outline for discrimi- ribbed, serpenticonic form with few sutural elements, which nation, but nothing more can be attempted with the mate- could be the Middle Devonian ancestral representative. rial that is available. Triainoceras is known from several species. The type Distribution.–Eastern North America, Europe, North species, by monotypy, is Goniatites costatus d’Archiac & de Africa, and Russia. It would seem that, on current evidence, Verneuil (1842: 340, pl. 31, figs 1, 1a) from Eibach and Schindewolfoceras and Wellsites are endemic to eastern North Oberscheld, Germany; the characters of the species were America. The near restriction of the family to siltstone facies worked out by Denckmann (1903b). T. gerassimovi Bo- in eastern North America is puzzling (Text-fig. 14). goslovsky (1958: 127, pl. 8, fig. 3; 1969: 279, pl. 25, fig. 3), Range.–European, North African, and Russian represen- from the Rudnyi Altai, Russia, is known in considerable on- tatives of the Triainoceratidae seem to be restricted to the togenetic detail. Uncertainly placed here (because sutures in Frasnian: Sandbergeroceras to Beloceras Genozones UD I-D- them are not known) are T.? incertum (d’Archiac & de H. Verneuil, 1842: 342, pl. 2, figs 6, 6a) from Brilon, Germany, and T.? tuberculosum (d’Archiac & de Verneuil, 1842: 342, Genus SANDBERGEROCERAS Hyatt, 1884 pl. 26, figs 4, 4a, 4b) from Oberscheld, Germany. Type species.–Sandbergeroceras sandbergerorum Miller, Miller (1938: 24, 178) grouped Sandbergeroceras, Schin- 1938, by subsequent designation (Miller, 1938: 178). dewolfoceras, and Pseudarietites as a new subfamily, the Sand- Diagnosis.–Triainoceratids with evolute inner costate bergeroceratinae. He considered Triainoceras as a synonym whorls and ribbing, often to 40 mm diameter, with smooth, of Sandbergeroceras and the same view has been put to us by laterally compressed outer whorls often approaching dis- R. T. Becker; this is likely, or even probable. But it cannot coidal form. Prominent growth-line lirae in inner whorls. be accepted until detailed ontogenies have been described Ribs prorsiradiate or approximately rectiradiate. Suture with of type material. It is now recognized that the Lower Car- a lobe centered on seam and four lateral lobes; ventral lobe HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 93

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

Specimen D WW WH UW WW/WH UW/D

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

deep and flat-bottomed with small siphuncular lobe within Sandbergeroceras syngonum Clarke, 1897 (Group) it; dorsal suture with mid-dorsal lobe and lobe between it Pl. 2, Figs 4-7, Pl. 3, Figs 5-6; Text-figs 27G-H; Table 3 and seam (full suture known only in the type species). Included species.–Sandbergeroceras sandbergerorum Miller Goniatites chemungense Clarke, 1885: 51. [1938: 178, pro Goniatites tuberculoso-costatus (pars) G. & F. Sandbergeroceras syngonum Clarke, 1897: 53; 1898, 1899a, b: 106- 108, ?text-figs 79-81, pl. 6, ?fig. 23, pl. 7, fig. ?19, fig. 20; Sandberger, 1850]; S. syngonum Clarke (1897, nom. nud.; House & Kirchgasser, 1993: 276. 1898, 1899a-b: 106, pl. 7, fig. 19; NYSM 4077, lectotype Sandbergeroceras? syngonum. Miller, 1938: 179-182, pl. 36, figs 9- here chosen); S.? enfieldense n. sp. (see below). 11. Discussion.–Sandbergeroceras was erected by Hyatt (1884: 333) with a misspelling as Sandbergeoceras (corrected by Type material.–Clarke illustrated and mentioned only Crosse & Fisher, 1884: 415; ICZN, 1956; see Miller, 1952). four specimens in describing this species. A specimen From the originally mentioned included species, Miller crushed flat on a bedding surface (first mentioned in 1885 named Goniatites tuberculoso-costatus as type species, but be- according to Clarke, 1898: 108, footnote) is here selected cause this was an invalid combination of two of d’Archiac & as lectotype (NYSM 4077, refigured here Pl. 3, Fig. 5). de Verneuil’s specific names, Miller (1938) proposed S. sand- Some doubt must surround the determination of the other bergerorum as a replacement name for certain figures of G. material that is assigned to the species with a question mark. & F.Sandberger (1850: pl. 4, fig. 1), which became the type There is a protoconch and initial whorl (NYSM 4078) and species. Because it is not clear that the figure refers to one a larger juvenile baritic replacement (NYSM 4075, refigured specimen (although it might), House & Ziegler (1977: pl. here, Pl. 2, Fig. 6) and a larger gutta percha mold referred to 6, figs 16-18) designated one specimen as lectotype and il- by Clarke as “probably” of the species (NYSM 4076, refig- lustrated it photographically. The genus differs from Triain- ured here, Pl. 2, Figs 4-5). These comprise the type series. oceras notably in the nature of the ventral lobe. Miller Additional material.–Only three additional specimens (1938), following Clarke (1897), but with a mark of doubt, are available, one collected by H. S. Williams from a locality referred S. syngonum from New York to the genus. in Tiogo County, south of Harford Mills (Cortland County; As applied to New York material, there is bound to be Loc. 64), preserved as a partial external mold (NYSM doubt surrounding the use of this generic name because no 12026, figured here Pl. 2, Fig. 7) and two loose specimens New York specimen referred here shows a complete suture. (NYSM 16577 and 16578) found by us at the same locality. All assignments are therefore tentative. Dimensions.–See Table 3. Distribution.–Eastern North America (Sonyea Group, Description.–Lectotype subevolute, but whorl section New York, and ?West Virginia) and Europe (Germany). In cannot be determined. Ribs number approximately 14 in New York, from the Sonyea Group (Middlesex and previous half whorl at 43 mm diameter; ribs approximately ?Cashaqua Shale) in Ontario County and equivalent level(s) rectiradiate but with slight flexure. One new specimen (Middlesex Shale and Cashaqua Shale) in Tioga County. (NYSM 12026) shows 15 ribs in previous half whorl at ap- The West Virginia record (Miller, 1938) remains uncon- proximately 28 mm diameter. For earlier whorls, gutta per- firmed. In Germany, only the type material of the Sand- cha mold (NYSM 4076, Pl. 2, Figs 4-5, Text-figs 27G-H) berger brothers exists from the Rhenish Schiefergebirge at a shows that early whorls have ribs prorsiradiate on flanks that level not documented. become almost rectiradiate by 19 mm diameter. Relation of Range.–Sandbergeroceras Genozone UD I-D to Probelo- ribs to growth-line lirae well shown by NYSM 4076 and by ceras Genozone UD I-E, based on New York evidence. Harford Mills specimen (NYSM 12026); both showing lirae conforming with course of ribs and covering shell both over ribs and between them with approximately 8-10 lirae from 94 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

Text-fig. 27. Sutures of Triainoceratidae and Beloceratidae of the late Devonian of various localities. A. Wellsites tynani House & 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 specimen from the Frasnian of Western Australia figured by Glenister (1958). D. Triainoceras gerassimovi Bogoslovsky, 1958; diagram illustrating the adult suture based on a specimen from the Rudny Altai, Russia, figured by Bogoslovsky (1969). E. Triainoceras costatum (d'Archiac & de Verneuil, 1842), the type species of the genus; suture reconstructed from the figure of the holotype from Oberscheld (Eibach), Germany. F. Sandbergeroceras sandbergerorum Miller, 1938; suture of the type species of the genus based on a specimen from Oberscheld (Königzug), Ger- many, figured by G. & F. Sandberger (1850). G-H. Sandbergeroceras syngonum Clarke, 1897; NYSM 4076, based on a specimen "from a concretion on Honeoye Lake" and possibly from the Cashaqua Shale. G. Growth lines, X 9. H. Whorl cross section, X 2.5. I. Schindewolfoceras chemungense (Vanuxem, 1842); NYSM 4073, suture (reversed for comparison) based on the holotype, at ca. 73 mm diameter from Rhinestreet Shale equivalents near Owego (Loc. 69), Tioga County, X 2. J. Schindewolfoceras aff. chemungense; SUI 40080, suture of a spec- imen from Hwy 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.

crest to crest. Discussion.–All of this material leaves much to be desired, Nothing need be added to the original description of the but a distinctive pattern of ribs and growth-line pattern pro- inner whorls given by Clarke (summarized by Miller, 1938) vides a means of distinguishing the species. Sandbergeroceras except to remark that their relationship to the larger speci- sandbergerorum differs in several respects from S. syngonum, mens is inferred only. No adult sutures are known. notably in the absence of marked ventrolateral furrows and HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 95

sharp ventrolateral shoulders at comparable diameters (cf., (NYSM 12028). House & Ziegler, 1977: pl. 6, fig. 16) and the narrower ven- Material.–Two large specimens recorded as from the tral sinus band; ribs in the German specimen number ap- “Enfield” Formation: the holotype (NYSM 12028) from the proximately 12 in the previous half whorl at 28 mm Hungerford (University) Quarry (Loc. 66), near Ithaca, and diameter and they seem more sharply defined than in S. syn- another specimen (a plastotype) (NYSM 12027; PRI 8742) gonum. from Bald Hill, Caroline (Tompkins County; Loc. 94). Both Distribution.–Middlesex Shale (Sonyea Group) in On- are preserved mostly as external molds in fine siltstone. tario or Yates county and Tioga County. Possibly in the Diagnosis.–Species of Sandbergeroceras in which ribbing Cashaqua Shale (Sonyea Group) in Ontario and Yates coun- is lost by approximately 20 mm diameter and ribs become ties. The record from Tygart Valley (Randolph County), less frequent just before their loss. Adult much larger than in West Virginia, is unconfirmed (Miller, 1938; House, 1978). any other known member of genus. The lectotype (NYSM 4077) is recorded as from the Dimensions.–NYSM 12028 (holotype), D = ca. 150.0 “lower black band” from Snyder’s Gully, Naples, a creek that mm, WH = ca. 68.0 mm, Wh = ca. 46.0 mm, UW = 48.0 crosses the Ontario-Yates county line near Woodville (On- mm, UW/D = 0.32; NYSM 12027 (plastotype), D = ca. tario County; Loc. 46c); the preservation and lithology con- 140.0 mm, UW = 39.0 mm, UW/D = 0.28. firm that it came from the Middlesex Shale. The following Description.–Inner whorls strongly costate with ribs specimens reported from the Portage or Naples beds could numbering 12 in previous whorl at umbilical width of 17 have come from the Cashaqua Shale: NYSM 4075, a baritic mm in holotype and 11 at same umbilical width in plasto- specimen from a concretion at Honeoye Lake (Ontario type. Ribs rectiradiate where best developed but in plasto- County); NYSM 4076, a gutta percha mold of a specimen type, as ribs are lost they are progressively more prorsiradiate from a crinoidal concretion at Naples (Ontario County); and follow course of growth lines that sweep forward from NYSM 4078, a baritic specimen from possibly the same seam but are more rectiradiate past the not-very-well- crinoidal horizon at Naples. Although the labels with marked umbilical shoulder. Adult whorls smooth but with NYSM 4076 and 4078 refer to their source as “crinoidal slight nodes replacing ribs after approximately 20 mm di- blocks,” which could be from the Cashaqua, there is some ameter. Adult laterally compressed, perhaps suboxyconic. suggestion of High Point Sandstone lithology (Clarke & The holotype is a phragmocone; the diameter of the Luther, 1904). Naples is the type locality for the High Point complete conch could have exceeded 250 mm. It shows part Sandstone, a unit higher up in the West Falls Group. of a suture at the near-maximum diameter, suggesting the The Harford Mills (Loc. 64) specimen (NYSM 12026) presence of approximately five or more lobes on the flanks. from Tioga County collected by H. S. Williams is probably The saddle is well rounded and the lobe deep and almost from an upper Middlesex Shale equivalent. In 1998, we parallel-sided with a pinched acute lobe. found two loose fragments of the species (NYSM 16577 and Discussion.–These giant specimens differ from all known 16578) in siltstone and silty mudrock within a section of members of the genus in their size and ribbing pattern. They black shale at the locality. share giant size with Wellsites known from rather farther Because this is the only distinctive form known from the south, but that genus is oxyconic and has a closed umbilicus Middlesex Shale, the Regional Zone of Sandbergeroceras syn- as nearly as can be determined. Even though the ventral su- gonum refers to this level and up to the entry of the distinc- ture is not known, the species is assigned, with question, to tive Cashaqua Shale faunas with the next zone fossil, Sandbergeroceras. Probeloceras lutheri. It could be that the range of S. syngonum Distribution.–Known only from the Enfield Formation continues into the next zone or higher, but that is uncon- near Ithaca (Tompkins County); horizon undetermined but firmed. probably from equivalents of the upper Cashaqua Shale Range.– Frasnian: Sandbergeroceras Genozone UD I-D. (Sonyea Group) and lower Rhinestreet Shale (West Falls Regional Zone of Sandbergeroceras syngonum (18). Group). The holotype is from the Hungerford (University) Quarry (Loc. 66), southeast of Ithaca, and the plastotype is Sandbergeroceras? enfieldense n. sp. from Bald Hill, Caroline (Loc. 94), southeast of Brookton- Pl. 2, Figs 1-2, Pl. 3, Figs 2-3 dale (Tompkins County). The innermost whorls described under Sandbergeroceras? syngonum might belong to this Sandbergeroceras? enfieldense Kirchgasser & House, 1981: 48, nom. species. nud.; House & Kirchgasser, 1993: 276. Range.–Frasnian: ?Probeloceras or Prochorites or Mesobe- Etymology.–Named for the Enfield Formation, near loceras Genozone(s) UD I-?E, or F, or G. ?Regional Zone of Ithaca (Tompkins County), the source of the holotype Probeloceras lutheri (19) or Prochorites alveolatus (20) or 96 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

Naplesites iynx (21). Wellsites williamsi (Wells, 1956) Text-fig. 27B Genus WELLSITES House & Kirchgasser, 1993 Type species.–Wellsites tynani House & Kirchgasser, 1993, Beloceras williamsi Wells, 1956b: 751, text-figs 2A, C, pl. 82, fig. by original designation. 4; House 1962: 258. Wellsites williamsi. House & Kirchgasser, 1993: 273, fig. 5B. Diagnosis.–Oxyconic triainoceratid without ribbing, ex- cept perhaps in early whorls, with involute conch. Primary Type material.–A single incomplete specimen preserved ventral lobe with single division and seven lateral lobes (in- as a partial internal mold is the holotype (NYSM/CU ferred to have developed as umbilical lobes) with all ele- 40023); only a small portion of the body chamber is pre- ments rather deep. served. Distribution.–Lower Rhinestreet Shale (West Falls Diagnosis.–Differing from only other species in genus, Group; Tompkins and Chemung counties). Wellsites tynani, in having longer sutural elements and ab- Range.–Frasnian: ?Mesobeloceras or Beloceras Genozone sence in them of swollen form and pinched terminations. UD I-?G, H. ?Regional Zone of Naplesites iynx (21a) or Description.–Holotype illustrated and described by Wells Wellsites tynani (21b). (1956b), so only brief comment is required. Largest whorl height seen is approximately 80 mm, with last septum being Wellsites tynani House & Kirchgasser, 1993 at approximately 74 mm diameter. Only portion of umbili- Pl. 1, Fig. 9; Text-fig. 27A cus preserved, but clearly open although small. Whorl sec- Wellsites tynani House & Kirchgasser 1993: 273, fig. 5A. tions seem clearly oxyconic. Sutures (Text-fig. 27B) show at least nine lateral lobes with second lateral deepest; suture Type material.–Holotype, SUI 42418. seems somewhat distorted but lobes subacute and saddles Diagnosis.–As for genus but with larger lateral lobes rounded. swollen and terminations pinched. Discussion.–With the removal of this species from Belo- Description.–Holotype a large crushed specimen showing ceras, there is now no evidence for the Beloceras in New York phragmocone and inferred oxyconic form at 190 mm diam- or North America. eter. With body chamber, it would have exceeded 250 mm Distribution.–Bald Mountain (Loc. 68) southeast of in diameter. Ventral suture with in excess of 10 lateral lobes Ithaca (Tompkins County). The level, recorded as approxi- of which second lateral is deepest; lateral lobes (U in origin) mately 150 ft (46 m) above the base of the Cayuta Shale, decrease in size toward umbilicus. Larger lobes showing suggests a horizon above the Moreland Shale, but in any case pinched terminations. Inner whorls not visible. Suture as il- an early Rhinestreet Shale (West Falls Group) equivalent. lustrated in Text-fig. 27A. Range.–Frasnian: ?Beloceras Genozone UD I-?H. ?Re- Discussion.–This form clearly belongs to the group gional Zone of Wellsites tynani (?21b). named Beloceras by Wells (1956b) but does not show the sigmoidal lobes and saddles of that genus; the swollen lobes Genus SCHINDEWOLFOCERAS Miller, 1938 and pinched terminations show that this specimen should be Type species.–Goniatites chemungensis Vanuxem, 1842, by referred to the Pharciceratoidea. It is presumed to be a tri- original designation. ainoceratid that has lost ribbing, either completely or shown Diagnosis.–Triainoceratids with evolute to subevolute only in the innermost whorls, which are not seen in the shells and ribbing normally to at least 80 mm diameter, with holotype. ribs usually rursiradiate in outer whorls and with ventrolat- Distribution.–Lower Rhinestreet Shale (West Falls eral furrows. Suture with short, squat, V-shaped ventral lobe Group; Chemung County). The holotype, collected by M. and approximately six lateral lobes decreasing in size toward C. Tynan, is recorded from spoil in a roadcut for US Rte. 17, umbilical seam. Dorsal suture unknown. 0.16 km east of the crossing of US Rte. 17 and Watercure Included species:–Schindewolfoceras chemungense (Vanux- Road, Elmira (Chemung County; Loc. 70), at an altitude em, 1842: 182, text-fig. 49, 1); S.? equicostatum Hall (1874: of approximately 910 ft (277 m). The horizon is said to be 3-4; 1876: pl. 69, fig. 10); S.? aff. equicostatum (Hall, 1874). the Moreland Shale, the basal tongue of the Rhinestreet Discussion:–Schindewolfoceras was proposed by Miller Shale, but see comments in locality description (Loc. 70). (1938: 181) with Goniatites chemungensis as the designated Range.–Frasnian: ?Mesobeloceras or Beloceras Genozone type species. He included in it the two named Chemung UD I-G or H. ?Regional Zone of Naplesites iynx (21a) or specimens of Vanuxem and Hall that were, at that time, re- Wellsites tynani (21b). ferred to Prolecanites. The two type specimens are not well preserved; only the holotype of S. chemungense shows a su- HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 97

ture and this was clearly a large goniatite (holotype, NYSM becoming nearly parallel-sided and subacute; ventral lobe 4073, figured herein, Pl. 2, Fig. 8, Text-fig. 27I). Both type broadly V-shaped, making first lateral saddle asymmetric. specimens were described by Miller (1938). The genus and Discussion.–No new material has come to light nor has species included in it are unknown outside New York State. anything comparable to this species been noted in other No additional material has been found in this study areas. For present purposes, a difference in rib frequency is apart from the specimen referred to Schindewolfoceras? aff. used to separate this species from Schindewolfoceras? equicostatum from Fairfield Forest (Loc. 67) near Speedsville equicostatum that has significantly more ribs at a comparable (Tioga County; NYSM 12027, Pl. 3, Fig. 1), and another diameter. Because the specimen is still septate, large adult but very poorly preserved specimen (NYSM 12025) from size, in excess of 200 mm diameter, is indicated. near Chemung (Chemung County; Loc. 95), figured here as Distribution.–Rhinestreet Shale equivalent (West Falls ?Schindewolfoceras sp. (Pl. 2, Fig. 3). Group; Tioga County). Vanuxem did not give a precise lo- Distribution.–West Falls Group in Tioga and Chemung cality for his specimen, but Hall (1879: pl. 69, fig. 9) refig- counties. Both type specimens are from the “Chemung” ured it and gave the locality as “Chemung near Owego, Group and the localities suggest West Falls Group equiva- N.Y.” This is our Loc. 69 (Tioga County). The horizon is lents. Schindewolfoceras? aff. equicostatum at Loc. 67 (Tioga thought to be an equivalent of the Rhinestreet Shale. Miller County) might be a mid-Rhinestreet occurrence. (1938: 184) gave the locality as “Cayuta Shale member Range.–Frasnian: ?Mesobeloceras to Beloceras Genozone (Spirifer disjunctus zone) of the Chemung Formation.” UD I-?G to ?H. Range.–Frasnian: ?Beloceras Genozone UD I-?H. Re- gional Zone of Schindewolfoceras chemungense (21c). Schindewolfoceras chemungense (Vanuxem, 1842) Pl. 2, Fig. 8; Text-fig. 27I Schindewolfoceras? equicostatum (Hall, 1874) Pl. 3, Fig. 4 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, Goniatites Chemungensis var. equicostatus Hall, 1874: 3-4; 1876, fig. 9, pl. 74, fig. 6; Hyatt, 1884: 333. pl. 69, fig. 10. Schindewolfoceras chemungense. Miller, 1938: 182-184, fig. 38B, Sandbergeroceras Chemungense Clarke, 1898, 1899a, b (pars): 108. pl. 36, figs 7-8; Bogoslovsky, 1969: 282-283; House & Kirch- Schindewolfoceras? equicostatum. Miller, 1938: 184, pl. 36, fig. 12; gasser, 1993: 276, 278. Bogoslovsky, 1969: 283; House & Kirchgasser, 1993: 276.

Type material.– The holoype (NYSM 4073), figured by Type material.–Only known by the holotype (NYSM Vanuxem & Hall, is poorly preserved in fine silstone; it is 4074) preserved in buff-weathering siltstone as a crushed in- laterally compressed and distorted and the body chamber is ternal and part external mold; incomplete and not showing not preserved. suture. Dimensions.–The holotype (NYSM 4073): D = ca. 78.0 Description.–Adult subevolute; whorls with well-rounded mm, Wh = ca. 28.0 mm, UW = ca. 31.0 mm, UW/D = convex flanks passing to rounded, weakly developed umbil- 0.40; S. aff. chemungense (SUI 40080, Pl. 4, Figs. 1-4): D = ical shoulder. Evidence of double ventrolateral furrow, but 41.2 mm, WW = 14.8 mm, WH = 15.4 mm, Wh = 10.3 venter not seen; maximum diameter seen approximately 70 mm, UW = 16.1 mm, WW/WH = 0.96, UW/D = 0.39. mm. Ribs well developed, forming and slightly prorsiradiate Description.–Holotype indicates rather evolute inner over umbilical shoulder, then passing slightly backward over whorls, becoming subevolute in outer whorls. Flanks appear flanks but lost after evidence of sharp forward projection to have been rounded and without marked umbilical shoul- into ventrolateral furrow. Ribbing almost completely lost in der; venter likely well-rounded but might have approached outermost quarter whorl (seen as external mold). Ribs num- subacute. No evidence of ventrolateral furrows but perhaps bering 14 in previous half whorl at 70 mm diameter. a feature of preservation. Ribs on inner whorls, at approxi- Discussion.–This form differs from Schindewolfoceras mately 22 mm diameter, 1.4 mm apart on mid-flanks, pro- chemungense in the greater frequency of ribbing and the ev- gressively becoming less prominent in last whorl seen and idence of ventrolateral furrows. In the absence of evidence of nearly rectiradiate on low flanks; earlier with backward de- sutures, the generic assignment must remain in doubt. flection and in last whorl with backward deflection on outer Distribution.–Unknown, but probably low in the West flanks. Ribs becoming weaker toward venter and numbering Falls Group of southern Tioga or Chemung counties. Orig- approximately nine in previous half whorl at ca. 70 mm di- inally said to come from Athens (Bradford County), Penn- ameter. Ventral sutures (Text-fig. 27I) comprising six lateral sylvania, and later said to come from a boulder, and hence lobes increasing regularly in size toward venter; outermost presumably loose. This could explain subsequent references 98 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

to it as from New York State because Athens is only a few Family ACANTHOCLYMENIIDAE Schindewolf, 1955 miles south of the state line near Waverley (Tioga County), This family is thought to have arisen from the Anarcestidae, and it might have been inferred to have been derived from particularly from the Atlantoceras group, by the division of the north. Hall (1879) considered that the brachiopods in the ventral lobe and the addition of other lobes between the the block indicatd that it came from the lower beds of the primary subumbilical lobe and the dorsal lobe. As presently Chemung Group which are well developed in the vinicity of conceived, the family comprises the genera Acanthoclymenia Waverly and Athens. It seems likely that the level is the early Hyatt, 1900, Pseudoprobeloceras Bensaïd, 1974, Ponticeras West Falls Group of modern terminology. Matern, 1929, Gogoceras Becker et al., 1993, Uchtites Bo- Range.–Frasnian: ?Mesobeloceras or Beloceras Genozone goslovsky, 1969, Prochorites Clausen, 1969 (= Eidoprobelo- UD I-?G or H. Regional Zone uncertain (21). ceras Kirchgasser, 1968, nom. nud.), and probably Enseites Becker & House, 1993, and a recent new genus, Chutoceras Schindewolfoceras? aff. equicostatum (Hall, 1874) Becker et al., 2000, based on material from the Timan-Pe- Pl. 3, Fig. 1 chora area of Russia. Acanthoclymenia, Ponticeras, Chutoceras, and Prochorites occur in North America. The family became Schindewolfoceras? aff. equicostatum. House & Kirchgasser, 1993: extinct with the Upper Kellwasser Event. 276. Distribution.–Eastern North America, Europe, Russia, North Africa, and Western Australia. Material.–A single specimen (NYSM 12027), preserved Range.–Upper Givetian to top of Frasnian: MD III-D in fine shelly siltstone, without any trace of sutures and in- to UD I-L. ferred to be a body chamber. Description.–Maximum diameter (estimated) approxi- Genus PONTICERAS Matern, 1929 mately 45 mm with convex flanks, prominent ventrolateral Type species.–Ammonites aequabile Beyrich (1837: 34) by groove with slight raised ridges on either side, and sugges- original designation. tions of tabular venter. Ribs numbering approximately 11 Diagnosis.–Conch evolute in early stages, subinvolute in in previous half whorl at ca. 45 mm diameter; these fading adult, usually laterally compressed, with phragmocone com- both in outermost flanks and toward aperture. Ribs with monly exceeding 100 mm diameter. Whorl section rounded slight backward deflection in earliest whorl seen but becom- to compressed. Venter tabular or rounded or oxyconic. With ing rather more rectiradiate; with evidence of forward sweep or without ventrolateral furrows. Suture as in Acanthocly- of ribs, as they are lost, toward ventrolateral groove. menia but some with divided ventral saddle very large. Lat- Remarks.–This is a very similar conch to Schindewolfo- eral saddle broadly arched, symmetrical to strongly ceras? equicostatum but differs in the rather earlier loss of rib- asymmetric with steep dorsal face; dorsal lobe deep. Lateral bing and in their lesser frequency. The latter is a lobe broadly to narrowly rounded, sometimes pointed. characteristic of S. chemungense, but the ventrolateral furrow Growth lines strongly to weakly biconvex; in some, is not (yet) known in that species. markedly rursiradiate across flanks with strong ventrolateral Distribution.–Middle Rhinestreet Shale equivalent (West salient. Falls Group; Tioga County). The single specimen collected Included species.–Ponticeras aequabile (Beyrich, 1837), P. by J. W. Wells is from Fairfield Forest (Loc. 67) near barroisi (Wedekind, 1917, 1918), P. perlatum (Hall, 1874, Speedsville (Tioga County), from the Upper Cayuta Shale at 1876), P.auritum (Holzapfel, 1899), P.bisulcatum (Keyser- approximately 400 ft (122 m) above a Tropidoleptus horizon. ling, 1844), P. discoidale Glenister, 1958, P. domanicense From a level between the Moreland Shale and Roricks Glen (Holzapfel, 1899), P. kayseri (Wedekind, 1917, 1918), and Shale and thus a mid-Rhinestreet Shale equivalent. P. keyserlingi (Holzapfel, 1899). Range.–Frasnian: Uncertain Genozone UD I-?G, H. Re- Discussion.–A neotype for Ponticeras aequabile was gional Zone uncertain (?21-22). erected by House & Ziegler (1977: 77) on the assumption that Beyrich’s specimen had been lost, it not having been Superfamily GEPHUROCERATOIDEA Frech, 1897 noted for a century and a half. The neotype was designated This major group, typical of the Frasnian and extinct by its as a specimen that had been figured by Wedekind (1918: pl. close, is thought to have arisen from simple serpenticonic 21, fig. 6) as the species (it was refigured by House & anarcestids. It includes the Acanthoclymeniidae, Koeneniti- Ziegler, 1977: pl. 1, fig. 25). R. T. Becker recently discovered dae, Gephuroceratidae, and Beloceratidae, all of which are Beyrich’s specimen in the Museum für Naturkunde, Berlin. well represented in the New York faunas. The superfamily The designation of the neotype therefore is invalidated. R. became extinct in the latest Frasnian Kellwasser Event. T. Becker is preparing a description of the specimen that one HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 99

Table 4. Biometric data for Ponticeras perlatum (Hall, 1874) from the Lodi Limestone. See Text-fig. 32 for graphs.

Specimen D WW WH UW WW/WH UW/D

NYSM 12166 ca. 92 ca. 15 ca. 34 ca. 32 0.44 0.35 ca. 63 ca. 11 ca. 25 ca. 22 0.44 0.35 ca. 45 ca. 9 ca. 16 ca. 16 0.56 0.36 NYSM 13173 60.8 11.3 24.3 19.3 0.47 0.32 41.5 9.1 17.2 12.4 0.53 0.30 28 ca. 6.0 11.9 7.7 0.50 0.28 8.2 2.5 3.5 2.8 0.71 0.34 5.6 1.9 2.0 2.2 0.95 0.39 4.3 1.4 1.4 1.7 1.00 0.40 3.4 ca. 1.1 1.1 1.3 1.00 0.38 2.5 ca. 0.90 0.83 0.90 1.08 0.36

of us (MRH) has examined. Beyrich’s original description and Truxton (Cortland County). We also include a few spec- and illustration are good and the specimen shows a whorl imens from more eastward localities collected by Charles section with a narrowly tabular venter. In this it differs from Thayer during a study of the paleoecology of the Genesee most New York specimens but agrees with the many, usually Group (Thayer, 1974). Most of the Thayer material is larger, specimens that Bogoslovsky (1969) described from poorly preserved molds but there is a fine specimen from a the Timan-Pechora region of northern European Russia. It tongue of black shale (Geneseo Shale) from near Upperville raises the question as to whether earlier members, such as P. (Chenango County; Thayer’s loc. A-9051) and two speci- perlatum, with rounded venters, should in the future be tax- mens from the Sherburne Siltstone from localties southeast onomically distinguished from a later group with tabular of Truxton (Cortland County; Locs A9157 and A9158). venters. Thayer’s collection will be deposited at the NYSM. Distribution.–Late Middle Devonian to mid-Frasnian. Dimensions.–See Table 4. MD III-D to UDI-E. Diagnosis.–Adult shell large, to 90 mm diameter, evolute and compressed, with flanks curving gradually to weak ven- Ponticeras perlatum (Hall, 1874) trolateral furrows and narrowly rounded venter. Adult suture Pl. 4, Figs 5-9, Pl. 5, Figs 1-9; with broadly rounded median saddle, deeply pointed ven- Text-figs 28A-G, 29A-C, 32A; Table 4 trolateral lobe, high, broadly rounded and slightly asymmet- ric lateral saddle, and broadly rounded lateral lobe. Growth Goniatites complanatus perlatus Hall, 1874: 1-2; 1875: 132-134; lines strongly sinuous in adult with prominent, rounded 1876: pl. 70, fig. 12; 1879: 458-459, pl. 70, fig. 12. salient, dorsad of ventrolateral furrow, which appear as low Goniatites complanatus (Hall, 1874). Hall, 1879 (in part): 455- 457, pl. 70, figs 6-8. arcuate ribs on internal molds (Pl. 5, Figs 8-9). Growth-line Manticoceras perlatum. Miller, 1938: 95-97, pl. 24, figs 11-12. lirae on flanks of inner whorls relatively straight and fascic- Ponticeras perlatum. Wells, 1956a: 748; House, 1962: 256, pl. 45, ulate, becoming more sinuose and evenly spaced. See also figs 10-12, text-figs 3A-B; Kirchgasser, 1975: 86-88, pl. 3, figs Kirchgasser (1975: 86-88). 6-11, text-fig. 11, table 4; House & Kirchgasser, 1993: 274- Discussion.–Ponticeras perlatum enters the New York sec- 276. tion near the top of the Geneseo Shale in the form of mostly crushed specimens at localities near Cayuga Lake (Tompkins Type specimen.–NYSM 3649, specimen figured by Hall County; Pl. 4, Fig. 6). Better specimens with preserved shells (1876: pl. 70, fig. 12; 1879: pl. 70, fig. 12) and herein (Pl. are in the main bed of the Lodi Limestone near the base of 4, Fig. 9) from the Sherburne Siltstone, Genesee Group, the overlying Sherburne Siltstone in the Seneca and Cayuga Homer (Cortland County). lake valleys, and especially from Lodi Glen (Loc. 27) near New material.–New specimens from the Lodi Limestone Lodi (Seneca County), and Romulus Town Quarry (Loc. at Romulus Town Quarry (Loc. 27a), Ovid, and one from 27a), near Ovid (Seneca County). Fragments of P.perlatum the Lodi at Beards Creek (Loc. 18/4) were added to the col- have been noted in the nodular westernmost bed of the Lodi lections in the NYSM and the CU made by various workers at Beards Creek (Loc. 18/4) in the Genesee Valley (Text-fig. over the past century from the Geneseo Shale, Lodi Lime- 28F). A specimen with a rather narrow lateral lobe from the stone and Sherburne Siltstone from localities between third cycle of the Lodi Limestone higher in Sherburne at Seneca and Cayuga lakes and eastward to Homer, Cortland, Seneca Lake (Loc. M, Ov-13) is referred to P. cf. perlatum 100 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

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 Sherburne Siltstone at Ithaca, Tompkins County (from House, 1962: text-fig. 3B), X 1.1. B, G. NYSM 13173, suture and cross section at 61.3 mm diameter, based on a specimen from the Lodi Limestone at Ovid (Loc. 27a, WTK Loc. 3012), Seneca County (from Kirchgasser, 1975), X 1.7. 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 the Lodi Limestone, Hubbard Quarry (Loc. 27b), southwest of Kidders, Seneca County. F. NYSM 16567 [3852/1], cross section based on a specimen 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, South Hill (Loc. AA, Dy-10c), Ithaca, Tompkins County, X 0.8. Scales = 5 mm (F); 1 cm (E, H-I). HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 101

Text-fig. 29. Sutures and cross sections of Ponticeras spp. from the Lodi Limestone (lower Penn Yan Shale) of New York State. A-C. Ponticeras perlatum (Hall, 1874). A. NYSM 12166, cross section of a specimen from the Lodi Limestone (Lodi B) at Romulus Town Quarry (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. 24 mm whorl height, based on a specimen from the Lodi Limestone (Lodi B) from the same locality, X 3.6. D-E. Ponticeras cf. perlatum, NYSM 12168, cross section and suture (reversed) at ca. 30 mm whorl height, of a specimen collected by G. Baird from the third cycle of the Lodi Limestone (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 Tommy Creek, 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). 102 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

(Text-figs 29D-E). and 3.4 m below the base of the Middlesex Shale, at Beards Ponticeras perlatum was collected or noted from a num- Creek (Loc. 18), near Leicester (Livingston County). ber of localities and horizons within the Sherburne Siltstone Discussion.–The shells are juvenile stages and are too in the Cayuga Valley by several workers beginning in the small to describe accurately. The preserved inner whorls are nineteenth century, most notably Williams (1884), Kindle evolute to serpenticonic and distinctly more rounded and (1896), and Boekenkamp (1963) (see locality discussions). depressed than those of specimens of Acanthoclymenia aff. Most specimens are poorly preserved and correlations with neapolitana from the Fossil Log Horizon, 3.4 m below the specific horizons within the Lodi and Penn Yan to the west base of the Middlesex. Previously this form was referred to have not been established (see Baird et al., 1989). Similarly Archoceras (Atlantoceras) sp. (Kirchgasser & House, 1981). the records of P.perlatum from Cortland County are poorly In Archoceras, however, lateral furrows are typically present. constrained stratigraphically, but all are presumably from There is some indication that the mid-ventral lobe is divid- the upper Geneseo Shale or Sherbrune Siltstone or equiva- ing. Consequently we designate the specimens as Ponticeras? lents. sp. It is now considered that this material cannot be referred Distribution.–Uppermost Geneseo and lower Penn Yan to Manticoceras evolutum (see House & Kirchgasser, 1993: shales (and equivalent Sherburne Siltstone to base of Ren- 276-277). wick Shale) (Genesee Group), from Livingston to Cortland Distribution.–Uppermost West River Shale (Genesee counties. Although Ponticeras can occur lower down in the Group; Livingston County). Geneseo Shale (Geneseo Limestone Horizon and Fir Tree Range.–Frasnian: Timanites Genozone UD I-C. Re- Pyrite), undoubted P.perlatum first occurs in the uppermost gional Zone of Koenenites beckeri (17b). Conodont transitans Geneseo Shale (Givetian conodont disparilis Zone) and Zone; MN Zone 4. ranges upward into the Lodi Limestone (Givetian conodont norrisi Zone) in the lower Penn Yan Shale and equivalent Genus ACANTHOCLYMENIA Hyatt, 1900 Sherburne Siltstone. In the Cayuga Valley, P.perlatum ranges Type species.–Clymenia (Cyrtoclymenia) Neapolitana to the top of the Sherburne Siltstone at its contact with the Clarke, 1892, by original designation (Hyatt, 1900: 548). overlying black Renwick Shale, beds that correlate westward Diagnosis.–Acanthoclymeniids in which Manticoceras- to horizons with lower Frasnian MN Zone 1 conodonts. At type suture is developed at least by 10 mm diameter. Conch Abbey Gulf (Loc. 21), in the Honeoye Lake Valley, Pon- subevolute, with flattened venter and ventrolateral furrows ticeras occurs in Penn Yan Black Shale B (above the Lodi and and ornamented in early five volutions with distinctive con- below the Renwick), a level in MN Zone 1 (Kirchgasser, cave lateral flares on flanks. Growth lines biconvex. 1994: fig. 4). Farther west, in the Genesee Valley, P.perlatum Included species.–Acanthoclymenia neapolitana (Clarke, can range higher in the Penn Yan to levels above the SB 1892: 57-63, text-figs 1-12; House, 1961: pl. 75, figs 1-11, black shale horizon (conodont MN 1 Zone) but probably text-figs 1A-D, lectotype NYSM 3625); A. genundewa does not overlap the lower range of Koenenites (conodont (Clarke, 1897: 53, nom. nud.; 1898, 1899a, b: pl. 8, figs 1- MN Zone 2). 3); A. forcipifer (G. & F. Sandberger, 1850: 81, pl. 6, figs Range.–Uppermost Givetian to lower Frasnian: lower 3a-c); A. orientale (Bogoslovsky, 1969: 89, text-fig. 19, pl. 2, Ponticeras or Petteroceras Genozone MD III-E to upper Pon- figs 3-5); A. planorbe (G. & F. Sandberger, 1850: 96, pl. 9, ticeras or Neopharciceras Genozone UD I-A. Regional Zone fig. 3a; lectotype of House in House & Pedder, 1963: 517). of P. perlatum (15b). Conodont disparilis and norrisi Zones Discussion.–The name Acanthoclymenia is employed here of Givetian to lower falsiovalis and MN Zone 1 of Frasnian for small, evolute to serpenticonic forms with tabular venters (Kirchgasser, 1994). in which the lateral saddle is high, asymmetrical, and some- what crooked but in which the saddles and lobes lack the Ponticeras? sp. extreme angularity of Probeloceras exemplified by the type P. Text-fig. 33M lutheri, here regarded as the primitive member of the Belo- ceratidae. Prochorites is separated from Acanthoclymenia and Archoceras (Atlantoceras) sp. Kirchgasser & House, 1981: 42-43. Probeloceras by its distinctively concave venter in the adult. Manticoceras cf. evolutum Petter, 1959. House & Kirchgasser, For 70 years, the type species (by monotypy) was consid- 1993: 276-277. ered to be a clymeniid. Clarke (1892) originally assigned it to the genus Clymenia and later Hyatt (1900: 548) created Material.–NYSM 12169, a calcitic replacement from the the genus Acanthoclymenia with it as the only species. Schin- Beards Creek Horizon (18a/5), and six calcitic replacements dewolf (1955: 422) placed the genus as the sole member of from the Fossil Log Horizon (18a/4b) (Williamsburgh Bed a new family, the Acanthoclymeniidae. As a member of the of Over et al., 1999, 2003) in the West River Shale, 2.4 m HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 103

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.

Specimen D WW WH UW WW/WH UW/D

NYSM 3632 12.4 ca. 3.5 5.2 4.8 0.67 0.39 NYSM 3631 8.0 ca. 2.6 2.8 3.4 0.93 0.43 NYSM 12171 ca. 7.7 2.4 2.3 — 1.04 — NYSM 3629 7.5 3.0 2.3 3.8 1.30 0.51 NYSM 3633 6.7 2.5 2.2 3.4 1.14 0.51 NYSM 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 —

Clymeniida, the form was anomalously old, as was recog- section as Acanthoclymenia sp. in the Penn Yan Shale just nized in the last century. In Europe, clymeniids do not ap- below the Crosby Sandstone at Keuka Lake but it can begin pear until the Platyclymenia Stufe of the Famennian. House slightly lower at the level of the Linden Horizon at Linden, (1961), however, was able to demonstrate from the baritic New York. A. genundewa ranges through the Genundewa type material that the supposed dorsal siphuncle was in fact Limestone and possibly into the lower West River Shale. the prolongation of a middorsal lobe and that the suture was From higher in the West River Shale, particularly in the also typical of Manticoceras. Acanthoclymenia and the Acan- rhythm below the Bluff Point Siltstone and in the Fossil Log thoclymeniidae were thus regarded as synonyms of Mantic- Horizon near the top of the unit, pyritic inner whorls of oceras and the Gephuroceratidae, respectively. rather more evolute forms occur that are referred to A. aff. In resurrecting the generic name for use as a genus neapolitana. Acanthoclymenia has not been found in the within the Gephuroceratoidea, we are following the reason- lower Cashaqua Shale but reappears in the upper Cashaqua able current trend to distinguish forms previously assigned as A. neapolitana, especially in the Shurtleff Septarian Hori- to the carpet-bag genus Manticoceras, but with distinctive zon. shell morphology, as distinct genera. The forms included in Range.–Frasnian: Upper Ponticeras or Neopharciceras Acanthoclymenia here all show rather more evolute shells Genozone UD I-A to Prochorites Genozone UD I-F. New than Probeloceras, and they differ also in that they achieve York: Koenenites Genozone UD I-B, Regional Zone of K. the Manticoceras type suture at an early stage, whereas most styliophilus styliophilus (16) to Prochorites Genozone UD I-F, species of Probeloceras never show it, and those that do Regional Zone of Prochorites alveolatus (20). achieve it late. It is considered that the plexus Ponticeras /Acanthoclymenia /Probeloceras is the line of evolution into Acanthoclymenia neapolitana (Clarke, 1892) the Beloceratidae. Pl. 6, Figs 1-15; Text-figs 30E-H, 32B; Table 5 The relationship of Ponticeras to the probeloceratid group and Manticoceras has been much discussed (Glenister, Clymenia (Cyrtoclymenia) Neapolitana Clarke, 1892: 57-63, text- 1958; Clausen, 1969; Bogoslovsky, 1969; Kirchgasser, figs 1-12; Frech, 1913: 8. Clarke, 1897: 54; 1898, 1899a, b: 131, 1975) but was not satisfactorily resolved until precise strati- Cyrtoclymenia Neapolitana. text-figs 102-106, pl. 8, figs 19-25. graphical information was forthcoming from sequences in Acanthoclymenia neapolitana. Hyatt, 1900: 548; Schindewolf, the Canning Basin (Becker et al., 1993). Most species of 1923: 482; 1934: 346; Miller, 1938: 189, figs 39A-I, pl. 38, Ponticeras are large in the adult compared to probeloceratids, figs 17-20; Schindewolf, 1955: 421; Schindewolf in Moore, with the problem in separation arising when comparing ju- 1957: L40, text-figs 39a-b; House & Kirchgasser, 1993: 276, veniles. Nevertheless we take the view that Ponticeras consti- 278. tutes a distinct group in which the external saddle and lobes Manticoceras neapolitanum. House, 1961: 472, pl. 75, figs 1-11, are rounded at all stages and the lateral saddle is low, sym- text-figs 1A-D; 1962: 258; Bogoslovsky, 1969: 222; Clausen, metrical, and broad. 1969: 116, 146; 1971: 182; Kirchgasser, 1975: 67. Distribution.–Eastern North America, Europe, Russia, Type material.–Miller (1938: 192) mentioned 21 syn- and Western Australia. New York: Penn Yan Shale (Genesee types (excluding one he regarded as a Manticoceras) and 29 Group) to Cashaqua Shale (Sonyea Group; Wyoming to hypotypes in the NYSM. These include the lectotype Steuben counties). In New York, Acanthoclymenia enters the (NYSM 3625) designated by House (1961: 474) and other 104 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

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 locality and 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 the Genundewa Limestone, Beard's Creek (Loc. 18/6a) X 18. J. NYSM 13166, suture at ca. 2 whorls, based on a specimen from the same locality 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 specimen from 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 and outline based on a specimen probably from the same locality and horizon; suture reversed, X 7. Scales = 1 mm (A); 5 mm (B). HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 105

material figured by Clarke [1899a, b: pl. 8, fig. 19 (NYSM are two pyritic and crushed specimens, one from the upper, 3631), fig. 20 (NYSM 3632), fig. 21 (NYSM 3628), fig. 22 dark transition shales of the Cashaqua Shale in Beards Creek (NYSM 3633), and fig. 25 (NYSM 3629). All of this mate- (Loc. 38/10; NYSM 16585), and the other from Buck Run rial is excellently preserved as baritic shell replacements. Creek (Livingston County; Loc. 39/22; NYSM 16586), but New material.–Nine indifferently preserved baritic re- these poorly preserved specimens could be Probeloceras or placements (including NYSM 12171 and 12172) and two Prochorites. crushed pyritic specimens, all collected by WTK. Range.–Frasnian: Prochorites Genozone UD I-F.Regional Dimensions.–See Table 5. Zone of Prochorites alveolatus (20). Conodont Zone MN 6. Description.–Detailed accounts of the type material hav- ing been given by Clarke (1899a, b), Miller (1938), and Acanthoclymenia aff. neapolitana (Clarke, 1892) House (1961), no lengthy account is required here. Text-fig. 30D Species characterized by unusual varices (festoons or flares) developed during the first five whorls on flanks and Material.—Fourteen calcitic and pyritic replacements, numbering 9-11 per whorl. Varices not extending onto the mostly small juveniles, from the Fossil Log Horizon of the venter or ventrolateral shoulder unless, as Clarke remarked, upper West River Shale (Williamsburgh Bed of Over et al., they did and were subsequently resorbed before formation of 1999, 2003), 3.4 m below the Middlesex Shale at Beards subsequent whorl. One specimen (Pl. 6, Fig. 1) suggesting Creek (Loc. 18a/4b), and three from probably the same accommodation of a later whorl to form of the flares; most horizon, 3.4 m below the Middlesex Shale at Chidsey Point specimens (Pl. 6, Figs. 9, 11, 13, 15) suggesting varices re- Creek (Loc. JJ, Py-7/2), Keuka Lake. stricted to the flanks. Discussion.–The small size and indifferent preservation Discussion.–It is the presence of lateral varices and a de- makes comparison of these specimens difficult. However pressed whorl section in the early volution that distinguishes they are all highly evolute to serpenticonic with a corre- this species from other widely evolute forms in the Cashaqua spondingly low rate of whorl expansion (see cross section, Shale. The holotype of Gephyroceras holzapfeli Clarke, 1898, Text-fig. 30D, NYSM 12148, from Beards Creek). Where is a pyritic internal mold, and hence does not show the shell seen, the lateral saddle is rather narrow and rounded. These (the festoonlike structures seen on the figure herein (Pl. 25, features suggest closer affinity to Acanthoclymenia neapoli- Fig. 1) represent areas of crushing within camarae of the tana from the Cashaqua Shale than to A. genundewa or A. cf. phragmocone), but it is a larger species and seems more evo- genundewa from the Genundewa Limestone and the Ge- lute at comparable diameters; this species is placed here as a nundewa Pyrite and lower West River Shale. junior synonym of Probeloceras lutheri. Enseites sulcatum Acanthoclymenia sp. A of Kirchgasser (1975) (NYSM (Matern, 1931), generically separated by Becker & House 13175) from the upper West River at Whetstone Brook (1993: 113), differs in a grooved venter shown by outer (Loc. 26a/1), 2 mi (3.2 km) west of Honeoye (Ontario whorls of the internal mold and in a rather more serpenti- County), differs from A. aff. neapolitana in its higher whorl conic form. expansion and broader lateral saddle. The stratigraphic po- Distribution.–Upper Cashaqua Shale (Sonyea Group) in sition of A. sp. A below the Middlesex Shale is unknown but Wyoming (Livingston and Ontario counties). The original is probably lower than the Fossil Log Horizon (Williams- syntypic material of Clarke almost certainly came only from burgh Bed) of Beards Creek and Chidsey Point (Text-fig. the Shurtleff Septarian Horizon in Shurtleff’s Gully (Loc. 9). 41) or Shurger’s Glen, a tributary of Kinney Creek, Livonia Distribution.–Fossil Log Horizon (Williamsburgh Bed) (Livingston County). This level is high in the Cashaqua of upper West River Shale (Genesee Group; Livingston and Shale (Text-fig. 10). Some of the material figured by Clarke Steuben counties). in 1898 and 1899, however, although almost certainly from Range.–Frasnian: Timanites Genozone UD I-C. Re- the same level, was recorded as from “calcareous concretions gional Zone of Koenenites beckeri (17b). Conodont Zone in the vicinity of Honeoye Lake (Ontario County) and to MN 4. the westward as far as Conesus Lake” (Clarke, 1899a, b: 134) and “various localities in Wyoming County” (Clarke, Acanthoclymenia genundewa (Clarke, 1898) 1899a, b: 159). Almost all of this material is in the NYSM. Pl. 24, Figs 1-4; Text-figs 30A-C, K, L; Table 6 New baritic material is also available from the Shurtleff Septarian Horizon at Shurtleff’s Gully (Loc. 41) (NYSM Gephyroceras? (Probeloceras?) genundewa Clarke, 1898, 1899a, b: 86, pl. 8, figs 1-3. 12171 and 12172; 16580-16583) and at Cottonwood Point Manticoceras genundewa (Clarke). Miller, 1938: 88-89, pl. 20, figs Gully (Loc. 40/13), Conesus Lake (NYSM 16584). There 1-4. 106 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

Table 6. Biometric data for Acanthoclymenia genundewa (Clarke, 1898) from the Genundewa Limestone.

Specimen D WW WH UW WW/WH UW/D

NYSM 13167 11.5 3.1 5.0 3.5 0.62 0.30 NYSM 16535 ca. 9.6 — ca. 5.0 2.9 — 0.30 5.1 1.5 1.9 2.1 0.79 0.41 3.7 ca. 1.1 1.3 1.6 0.85 0.43 2.7 ca. 9.0 0.90 1.1 1.00 0.41 NYSM 13162 9.1 — 4.3 2.5 — 0.27 NYSM 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 0.40 4.0 1.3 1.4 1.7 0.93 0.43 2.9 1.0 0.95 1.2 1.05 0.41 NYSM 13169 4.4 1.3 1.5 1.8 0.87 0.41 3.2 0.95 1.1 1.4 0.86 0.44

Probeloceras genundewa (Clarke). House, 1962: 256-259, text-fig. asymmetrical lateral saddle and broad, rounded lateral lobe; 3E; Kirchgasser, 1975: 77-82, pl. 2, figs 1-6, pl. 3, figs 3-4, greater asymmetry and angularity of lateral lobe is apparent 12-13, text-figs 7B, 8B, 9A-F, table 2. at approximately 6.5 mm diameter (NYSM 13162; Kirch- gasser, 1975: text-fig. 9E). Type material.—Clarke’s three syntypes are juveniles, the Distribution.–Lower Genundewa Limestone (Genesee largest (NYSM 3647) approximately 8 mm in diameter. The Group; Livingston, Ontario, and Yates counties) and prob- lectotype, selected herein, is NYSM 3645 (Clarke, 1898: pl. ably Harrell Shale in West Virginia. In the Genesee Valley 8, fig. 1; Miller, 1938: pl. 20, fig. 2; refigured here, Pl. 24, (Livingston County) and near Canandaigua Lake in Ontario Fig. 4) from the Genundewa Limestone at Genundewa and Yates counties, Acanthoclymenia genundewa appears to Point (Loc. 23b), Canandaigua Lake (Yates County). NYSM be restricted to the nodular lower part of the Genundewa 3646 (Pl. 24, Figs 1-2) is from the same horizon and locality, Limestone, below the well-bedded layers that define the but NYSM 3647 could have come from the Genundewa upper part of the unit. Limestone at Bristol (Ontario County; Miller, 1938: 89). From the Harrell Shale south of Landes Post Office Material.–New collections from the Genundewa Lime- (Grant County), West Virginia, House (1978) assigned to stone are: Taunton Gully (Loc. 17/8), near Leicester (Liv- Probeloceras cf. genundewa a collection of 16 specimens in ingston County), Beards Creek, Leicester (Livingston which whorl form and cross section conformed closely with County; Loc. 18/6a), Fall Brook-Dewey Hill, Geneseo (Liv- Acanthoclymenia genundewa from the Genundewa Lime- ingston County; Loc. 19/10), Seneca Point (Loc. 23/1), and stone in New York. Genundewa Point (Loc. 23b/4), Canandaigua Lake (On- The stratigraphic position of the single specimen from tario and Yates counties). Mostly calcitic replacements; some the Chut River section in Timan, Russia, described and il- baritic and pyritic replacements. Kirchgasser (1975) re- lustrated by House & Becker in Becker et al. (2000) as Acan- ported on specimens from the Genundewa Limestone at thoclymenia aff. genundewa, is unknown. Beards Creek (Loc. 18, WTK 3001) and at Seneca Point Range.–Frasnian: Koenenites Genozone UD I-B. Re- (Loc. 23, WTK 3006a). gional Zone of K. styliophilus kilfoylei (16). Conodont Zone Description.–Little can be added to Clarke’s (1898) de- MN 2, ?3. scription, summarized by Miller (1938), and to the account given by Kirchgasser (1975). Conch thinly discoidal and Acanthoclymenia cf. genundewa (Clarke, 1898) subevolute with whorl height appearing to increase more rapidly than in Acanthoclymenia aff. neapolitana from the Remarks.–At the NY Rte. 20 embankment (Loc. 24) and upper West River Shale and A. neapolitana from the upper beneath the bridge overpass at Bethany Center (Genesee Cashaqua Shale. Flask-shaped in cross section with rounded County; Loc. 24/2), several pyritic replacements of juveniles flanks converging to weak ventrolateral furrows and tabular from the basal West River Shale have whorl forms and su- venter. Growth lines with strong deep saddle on the mid- tures resembling Acanthoclymenia genundewa but better ma- flank. terial is needed for an unqualified assignment. Similarly Suture of the lectotype (NYSM 3645) at 4.8 mm diam- problematic are several calcitic replacements from the lower eter, illustrated by House (1962: text-fig. 3E), with broad HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 107

few meters of the West River Shale at Beards Creek, Leicester Included species.–The type species, Chutoceras manticoides (Livingston County; Loc. 18a/1-3). Becker & House (in Becker et al., 2000: 83, text-figs 8A-B, Also included in Acanthoclymenia cf. genundewa are sev- pl. 1, figs 1-2) and C. nundaium (Hall, 1874), described eral pyritic replacements of juveniles from the Genundewa herein. Pyrite at Keuka Lake, including one from Cornwall Gully Discussion.–The distinguishing feature is the ponticeratid (Loc. DD, Py-10/1), Penn Yan (Yates County), and several form with a suture of the complexity of Koenenites, that is from Sunset Point Gully (Loc. F, Py-16/2), 9.3 km (5.8 mi) more advanced even than Manticoceras in possessing an um- southwest of Penn Yan (Yates County). bilical lobe centered on or near the seam. The shell form in- Distribution.–Genundewa Pyrite Horizon and basal and dicates relation to Ponticeras. The type species shows a lower West River Shale (Genesee Group) in Genesee (Liv- twisted first lateral lobe, a larger lateral saddle and sharp lat- ingston and Yates counties). eral lobe, and has the characteristic umbilical lobe probably Range.–Frasnian: Timanites Genozone UD I-C. Re- just dorsad of the umbilical seam. gional Zone of Manticoceras contractum (17a). Conodont Distribution.–Eastern North America: Ithaca Shale and Zone MN 3. Sandstone (Genesee Group) in Tompkins County; Timan, Russia. Acanthoclymenia sp. Range.–Ponticeras or Neopharciceras Genozone UD I-A (New York): ?Sandbergeroceras Genozone UD I-D, ?Probelo- Material.–Single pyritic inner whorl collected by J. Kral- ceras Genozone UD I-E (Timan, Russia). ick from a concretion in the Penn Yan Shale in the interval 0.30 m (1 ft) below the base of the Crosby Sandstone at Chutoceras nundaium (Hall, 1874) Sunset Point, Keuka Lake (Yates County; Loc. F, Py-16). Pl. 9, Fig. 1; Text-figs 28H-K Small crushed molds of an evolute goniatite from the shale on top of the Linden Horizon at Linden (Genesee County; Goniatites (Clymenia?) nundaia Hall, 1874: 3 (134-135); 1876: pl. Loc. 15/8), are indeterminate but could be Acanthoclymenia 70, figs 13, 15 (not fig. 14 = Ponticeras perlatum) Hall, 1879: 460-463, pl. 70, figs 13, 15, (not sp. Goniatites sinuosus fig. 14 = P.perlatum), pl. 74, fig. 11; 1888: 40, pl. 128, figs 1- Description.–Specimen NYSM 12170 with diameter of 2. 9.0 mm. Coil moderately evolute with high whorl expan- Ponticeras cf. regale (Holzapfel, 1899). Kirchgasser, 1985: 231-232, sion. Whorl width highest near umbilical wall; flanks slop- fig. 4b; House & Kirchgasser, 1993: 275-276. ing gently to narrow tabular venter; distinct but shallow ventrolateral furrow. Suture not seen but growth-line pattern Material.–Lectotype here designated as NYSM 3751, well displayed and showing deep saddle across flank and figured by Hall (1879: pl. 70, fig. 15, pl. 74, fig. 11). There strong, narrow, ventrolateral salient typical of Acanthocly- are four specimens from the Ithaca Shale and Sandstone menia. housed in the NYSM that are assigned to this species: Discussion.–Close affinity to Acanthoclymenia genundewa NYSM 3749 and 3751, originally described by Hall (1874, from the Genundewa Limestone is suggested. a valid designation) as Goniatites nundaia and later figured Distribution:–Upper Penn Yan Shale (Genesee Group) and described (Hall 1876, 1879) as Goniatites sinuosus. The in Yates County; ?Genesee County. localities given by Hall were “Lower Chemung group or Range.–Frasnian: Koenenites Genozone UD I-B. Re- Ithaca Beds of the Chemung Group . . . at the inclined-plane gional Zone of K. styliophilus styliophilus (16). Conodont of the railroad” at or near Ithaca (Tompkins County). The Zone MN 2. inclined plane locality (NYSM 3749 and 3751) is at South Hill, Ithaca (Loc. AA, Dy-10c), and the horizon is in the Genus CHUTOCERAS Becker & House, 2000 Ithaca Shale and Sandstone at approximately the level of the Type species.—Chutoceras manticoides Becker & House, University Quarry at Fall Creek, Ithaca (see below). A third 2000, by original designation (Becker & House in Becker et specimen (NYSM 3750) figured by Hall (1876: pl. 70, fig. al., 2000: 83). 14; 1879: pl. 70, fig. 14) from “the Ithaca beds of the Diagnosis.–Large laterally compressed acanthoclymenids, Chemung group near Truxton, New York (Cortland with wide umbilicus, similar to Ponticeras but not develop- County)“ is believed to be a Chutoceras but it is too poorly ing tabular venter and with more advanced sutures compris- preserved for certainty. ing sharp lobes and rounded saddles, often rather equal and Two additional specimens (NYSM 3753 and 3754) were distinguished by sharp umbilical lobe centered on or just described and figured by Hall (1888: 40, pl. 128, figs 1-2) dorsal of seam. as Goniatites sinuosus from the “Portage group. Ithaca, 108 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

Tompkins County.” Labels with the specimens and the mu- ployed for specimens subsequently discovered in what are seum catalog (Clarke & Reudemann, 1903: 605) indicate now the Genesee, Sonyea, and West Falls Groups (Hall, that they were collected by S. G. Williams from the “Portage 1876, 1879), Hall introduced the names Goniatites (Cly- (Ithaca beds) at the University quarry,” Ithaca, New York. menia?) nundaia, G. pattersoni (Hall, 1860), and G. com- There were many quarries on and in the vinicity of the Cor- planatus perlatus to differentiate some forms as species nell University campus during the latter part of the nine- separate from G. sinuosus. Both Clarke (1898, 1899a, b) and teenth century, but the one called the University Quarry was Miller (1938: 107) concluded that G. pattersoni is a syn- located at the edge of the Fall Creek (Loc. X, Dy-10) and the onym of Manticoceras sinuosum, an opinion with which we interval of shelly limestone horizons in the Ithaca Shale and agree. Similarly, Hall (1879) and Clarke (1898, 1899a, b) Sandstone (“Firestone bed” facies of Williams, 1884: 17) concluded that the type specimens of G. (C.?) nundaia were that yielded the specimens is estimated to be approximately also conspecific with the type of G. sinuosus and Miller 72 m (235 ft) above the base of the Renwick Shale (Kirch- (1938: 106-107) concurred. The figured types of G. (C.?) gasser, 1985). nundaia (NYSM 3749 and 3751), however, which are here Some additional but poorly preserved specimens referred to Chutoceras nundaium, are clearly not M. sinuo- (crushed molds) from the Ithaca Shale and Sandstone in the sum, a species characteristic of the Cashaqua Shale of the Ithaca area are here assigned to Chutoceras with question: Sonyea Group. CU 10928 and 10938 from approximately the level of the The name “nundaia“ is available for NYSM 3749 and Williams Brook Coquinite, approximately 53 m (175 ft) 3751. The first of these is strongly distorted and could be a above the Renwick Shale at Williams Brook (Loc. R, I-2); Ponticeras because it has a rounded lateral lobe and no lobe CU 10943, from a quarry in the lower Ithaca, 0.5 mi (0.8 on or near the umbilical seam. The two specimens (NYSM km) south of Williams Brook (Loc. S, I-2a); an unlabeled 3753 and 3754) assigned to Goniatites sinuosus by Hall CU specimen from the Ithaca in Enfield Glen (Loc. BB, I- (1888) are here assigned to Chutoceras nundaium. 8a); and CU 10923a from the Ithaca at Cascadilla Creek Hall’s specific name, which has clear priority, might be (Loc. Z, Dy-10b). A fragment with growth lines from the derived from “Nundawao,” the “Great Hill people” of the upper Penn Yan Shale at Mill Creek (Loc. L, Bed Ov-10/2) Iroquois whose settlement by the same name, said to be the near Lodi (Seneca County), could represent C. nundaium. oldest Seneca village, was located just north of Naples at the Dimensions.–NYSM 3753, D = 110 mm, WH = 43.7 southern end of Canandaigua Lake (Luther, 1910). The mm, UW = 32.8 mm. group’s first council fire took place to the north at “Genun- Diagnosis.–Large, evolute, widely umbilicate, com- dewa” or “Bare Hill” on the eastern side of the lake, an area pressed shells; diameter of NYSM 3753 exceeding 100 mm. that would link the name appropriately to rocks of the Whorl-form similar to that of Ponticeras perlatum but flanks Genesee Group. However, the name also suggests the area of more parallel-sided, with venter broader and flatter, and the Nunda Sandstone, a unit in the West Falls Group, with sharper break at ventrolateral margin. Weak ventrolat- named for Nunda (Livingston County), which is perhaps eral furrow as in P. perlatum not seen. confusing for specimens restricted to the stratigraphically Suture characterized by broad, prominent ventrolateral, much lower Genesee Group in Tompkins County, but Hall’s lateral, and umbilical saddles that are more symmetrical and name has clear priority. equally sized than in Ponticeras perlatum. Lateral lobes nar- We have referred this group in the past to Ponticeras and rower, more angular, and pointed than in P.perlatum. Chuto- to the species P. cf. regale (see Kirchgasser & House, 1981; ceras manticoides differs in having umbilical lobe not Kirchgasser, 1985; House & Kirchgasser, 1993) to draw at- centered on seam and slightly twisted, sharp first lateral lobe. tention to their similarity to one member of a group of pon- Growth lines follow pattern seen in Ponticeras perlatum ticeratids described by Holzapfel (1899) from the Lower with broad lobe centered on mid-flank leading to broad and Domanik Suite or Group, at localities along the Chut River, prominent salient centered outside ventrolateral margin, and near Ukhta, in South Timan, Russia. These Timan species, tongue-like lobe across venter. redescribed by Bogoslovsky (1969), are similar to the New Discussion.–When James Hall described the species Go- York specimens in their relatively large evolute shells and niatites sinuosus in 1843, only a few fragments were known, prominent growth lines with strong ventrolateral salients, one of which is the syntype [Hall, 1843: 243, fig. 106(6)] but do not show the sutures with broadly rounded, symmet- from Cashaqua Creek (Cashaqua Shale of the Sonyea Group rical lateral saddles, sharp lateral lobes. or umbilical lobes at Cashaqua Creek, Sonyea, Livingston County); the spec- centred on the seams. The Lower Domanik Ponticeras imen is AMNH 5887/1 (Miller, 1938: 115). species and Chutoceras manticoides are reported at levels with Although the name Goniatites sinuosus was widely em- conodonts of Zone MN 5 (punctata Zone) that is much HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 109

Table 7. Biometric data for Prochorites alveolatus (Glenister, 1958) from the upper Cashaqua Shale. See Text-fig. 32 for graph.

Specimen D WW WH UW WW/WH UW/D

NYSM 13159 19.9 — 8.9 5.5 — 0.28 NYSM 4063 17.5 4.1 8.0 4.7 0.51 0.27 NYSM 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 0.32 7.4 2.0 2.9 2.7 0.69 0.36 2.1 ca. 0.65 0.65 0.82 1.00 0.39 NYSM 13157 ca. 16.7 — ca. 7.5 ca. 4.4 — 0.26 NYSM 4064 15.6 4.2 7.6 3.8 0.55 0.24 NYSM 12747 4.6 — 1.8 1.8 — 0.38

higher than Ponticeras and Chutoceras in New York (Becker figs 1-5 (not figs 6-10 = P. lutheri), text-figs 70-71, 73-74, 77 et al., 2000). (not text-figs 68-69, 72, 75-76 = P. lutheri). Among the crushed molds from lower Ithaca Sandstone Probeloceras lutheri (Clarke). Miller, 1938 (pars): 63-70, figs 10C- and Shale that are assigned with question to Chutoceras 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). nundaium, CU 10938 from Williams Brook and the unla- Probeloceras alveolatum Glenister, 1958: 69-70, pl. 15, figs 2-3, beled specimen from Enfield Glen, in particular, show clear text-figs 3A-B; House, 1962: pl. 45, figs 5-6, text-figs 3C-D. evidence of the sharp break at the ventrolateral margin that Eidoprobeloceras strix Kirchgasser, 1968: 4175, nom. nud. is characteristic of the species. Probeloceras strix Kirchgasser,1975: 83-85, pl. 1, figs 6-12, 15, pl. Distribution.–Ithaca Shale and Sandstone (Genesee 3, figs 14-20, text-figs 6, 7B, 8B, 10, table 3. Group) in Tompkins County; ?upper Penn Yan Shale (Gene- Prochorites alveolatus. House & Kirchgasser, 1993: 276. see Group) in Seneca County. Chutoceras nundaium is re- stricted to the Ithaca Shale and Sandstone of the Genesee Type specimen.–By monotypy, BMR 1727 from 187-213 Group in Tompkins County, in the interval between the ft (57-65 m) above the base of the Gogo Formation, Locality Williams Brook Coquinite (and possibly lower in the Ithaca) K Dup 126, approximately 4 mi southwest of Old Bohemia and the Quarry Sandstone interval at Ithaca (Kirchgasser, Homestead (126-06E;18-42S), Western Australia. Specimen 1985). The species could be represented in the equivalent described by Glenister (1958). upper Penn Yan Shale in Seneca County. Material.–New York material was listed by Kirchgasser Range.–Frasnian: Ponticeras or Neopharciceras Genozone (1975). In addition there is a fine small specimen acces- UD I-A. Regional Zone of Chutoceras nundaium (15c). sioned under NYSM 3760 (12306/12). Conodont Zone MN 1 (Kirchgasser, 1994; conodonts re- Dimensions:–See Table 7 and Text-fig. 32C for graphs. covered from limestone matrix with NYSM 3753). Description.–A full description of New York material under the name Probeloceras strix was given by Kirchgasser Genus PROCHORITES Clausen, 1969 (1975). Type species.–Probeloceras alveolatum (Glenister, 1958), Remarks.–Kirchgasser (1975) removed from Probeloceras by original designation (Clausen 1969: 116). lutheri six of Clarke’s syntypes (NYSM 4061-4064, 4070, Diagnosis.–Acanthoclymenids similar to Acanthoclymenia and 4071) and redescribed them with 11 additional speci- but with grooved venter on mold and bicarinate venter on mens as P. strix. The name “strix” referred to the distinctive conch; suture as for family but with narrower lateral saddle concave venter in contrast to the tabular venter in P.lutheri. with sigmoidal ventrad face. Growth lines prorsiradiate with The striking similarity of the New York form with the then shallow ventral sinus and broad salient. single known specimen of Prochorites alveolatus from the the Distribution.–Eastern North America (Sonyea Group in Canning Basin of Australia was noted (Kirchgasser, 1975: New York) and Canning Basin, Western Australia. 85) and subsequent discoveries of the form in Australia have Range.–Frasnian: Prochorites Genozone UD I-F. Con- demonstrated their identity (Becker et al., 1993). odont Zone MN 6, 7. Distribution.–Upper Cashaqua Shale (Sonyea Group) in Livingston and Ontario counties, and Canning Basin, West- Prochorites alveolatus (Glenister, 1958) ern Australia. Pl. 7, Figs 4-6; Text-figs 31, 32C; Table 7 Range.–Frasnian: Prochorites Genozone UD I-F.Interna- tional and Regional Zone of Prochorites alveolatus (20). Con- Probeloceras lutheri (Clarke). Clarke, 1898 (pars): 89-102, pl. 7, odont Zone MN 6. 110 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

Text-fig. 31. Sutures and cross sections of Prochorites alveolatus (Glenister, 1958) from the Cashaqua Shale of New York State. A. NYSM 4063, 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 Naples Beds, Honeoye Lake, and probably from the Shurtleff Septarian Horizon, Livingston County (from House, 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 Shurtleff Septarian Horizon, Whetstone Brook (Loc.42/WTK loc. 3004), Honeoye, Ontario County, X 5.6. E. NYSM 13159, reversed suture at 6.00 mm 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 give either 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 thought to be from the Shurtleff Septarian Horizon, Whetstone Brook (Loc.42/WTK loc. 3004), Honeoye, Ontario County, X 5.6.

Prochorites aff. alveolatus (Glenister, 1958) deep umbilicus suggest affinity with Prochorites alveolatus from the upper Cashaqua Shale. Probeloceras aff. strix Kirchgasser 1975: 85, pl. 3, fig. 5. Distribution.–Lower Rhinestreet Shale (West Falls Group) in Livingston County. Remarks.—A single calcitic replacement (NYSM 13176) Range.–Frasnian: ?Mesobeloceras Genozone UD I-?G. Re- is known from the Fossil Log Horizon, 12.8 m (42 ft) above gional Zone Uncertain (?21). Conodont Zone MN 7. the base of the Rhinestreet Shale at Buck Run Creek (Loc. 65/1), Mt. Morris (Livingston County). Only the side of Family KOENENITIDAE Becker & House, 1993 the specimen is seen; the venter and suture are not The family Koenenitidae comprises two stocks, one contain- preserved. The moderately evolute coil, thinly discoidal ing Koenenites Wedekind, 1913, Hoeninghausia Gürich, whorl form with high whorl expansion and the relatively OS ICGSE:LT EOINGONIATITES DEVONIAN LATE KIRCHGASSER: & HOUSE

Text-fig. 32. Graphs showing the ontogeny of Ponticeras, Acanthoclymenia, and Prochorites spp., based on data given in Tables 4, 5, and 7. A. Ponticeras perlatum (Hall, 1874) from the Lodi Limestone. B. Acanthoclymenia neapolitana (Clarke, 1892) from the Shurtleff Septarian Horizon of the upper Cashaqua Shale. C. Prochorites alveolatus (Glenister, 1958) from the upper Cashaqua Shale. 111 112 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

Table 8. Biometric data for Koenenites styliophilus styliophilus (Clarke, 1898) from the Penn Yan Shale. See Text-fig. 35 A for graph.

Specimen D WW WH UW WW/WH UW/D

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 0.24 ca. 7.9 c. 2.9 ca. 3.4 2.0 0.85 0.25 ca. 5.3 2.2 2.6 1.4 0.85 0.26 3.4 1.7 ca. 1.3 1.2 1.31 0.35 2.6 1.2 0.9 0.8 1.33 0.31 1.9 0.95 0.68 0.73 1.40 0.38 NYSM 12140 ca. 55 — 25.9 13.5 — 0.25 NYSM 12139 ca. 38 10.2 18.1 — 0.56 — NYSM 12158 ca. 36 ca. 10.2 17.5 8.6 0.58 0.24 NYSM 12159 28.5 13.3 8.5 6.9 0.64 0.24 NYSM 12137 16 6.0 7.8 4.8 0.77 0.30 10.3 4.3 4.9 2.4 0.88 0.23 NYSM 12138 10.1 4.6 4.6 2.6 1.00 0.26 NYSM 12160 6.9 2.9 3.1 1.8 0.94 0.26 4.5 2.1 1.9 1.4 1.11 0.31 3.1 1.4 1.3 1.0 1.08 0.32 2.2 1.1 0.83 0.7 1.33 0.32 1.6 0.75 0.63 0.45 1.19 0.28 1.1 0.65 0.50 0.43 1.30 0.39 NYSM 12161 6.8 3.1 3.1 1.8 1.00 0.26 4.6 2.2 2.0 1.3 1.10 0.28 3.2 1.6 1.3 1.1 1.23 0.34 2.3 — 0.85 0.77 — 0.33 1.7 0.9 0.63 0.53 1.43 0.31

1896: 348, and Komioceras Bogoslovsky, 1958: 115, and the berger, 1850), K. lamellosus kirchgasseri House, 1978, K. other stock including Timanites Mojsisovics, 1882. Only the cooperi Miller, 1938, K.? fasciculatus (Clarke, 1898), K. hoen- first is represented in New York, but Timanites ocurs in inghausia (von Buch, 1832), K. styliophilus (Clarke, 1898), Western Canada (Miller, 1938). and K. beckeri n. sp. Distribution.–North America, Europe, Russia, and West- Discussion.–Early representatives have broad whorl forms ern Australia. with flanks sloping steeply to a narrowly rounded venter, Range.–Frasnian: Koenenites Genozone UD I-B to low, rounded saddles and lobes, and widely spaced growth- Timanites Genozone UD I-C, ?Beloceras Genozone I-H. line lirae. By comparison, later forms have more compressed, flat-sided whorls with broadly rounded to flattened venters, Genus KOENENITES Wedekind, 1913 sutures with high, asymmetrical lateral saddles, narrowly Type species.–Goniatites lamellosus G. & F. Sandberger, rounded to angular lobes, and more closely spaced growth- 1851: 85; 1850: pl. 8, fig. 1, by subsequent designation line lirae. (Wedekind, 1917: 126). Distribution.–North America (Squaw Bay Limestone, Diagnosis.–Conch subdiscoidal to stout, moderately evo- Michigan; Genesee Group, New York; Harrell Shale, Penn- lute, with ventrolateral furrows and rounded to flattened sylvania and West Virginia), Europe, and Western Australia. venter. External suture similar to that of Manticoceras but The earliest representatives of Koenenites in New York are K. with lateral saddle less prominent and with additional um- styliophilus styliophilus from the upper Penn Yan Shale and K. bilical lobe (U2). Ventrolateral lobe (E1) developing on styliophilus kilfoylei n. ssp. from the lower Genundewa Lime- flanks of external or ventral lobe following pattern of phar- stone, both closely similar to K. cooperi from the Squaw Bay ciceratids. Growth lines characterized by raised lirae that are Limestone of Michigan. Koenenites sp. of House (1978) and prominently developed, particularly as a salient on margins K. lamellosus kirchgasseri from the Harrell Shale of West Vir- of ventrolateral furrows. Ventral wrinkle layer consisting of ginia occur at a level equivalent to the Genundewa Lime- discontinuous, wavy, closely spaced lines. Hoeninghausia stone of New York. Also assigned to Koenenites, but with similar but developing oxyconic venter. question, is K.? fasciculatus from the lower Genundewa Included species.–Koenenites lamellosus (G. & F. Sand- Limestone and K.? cf. fasciculatus from the middle and upper HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 113

West River Shale. The last representatives of the genus in fig. 34E). These somewhat more massive shells have only New York are specimens assigned to K. beckeri n. sp. from been seen in oblique cross section and better preserved ma- the middle and upper West River Shale, which most closely terial will be needed to determine whether they are a sepa- resemble K. lamellosus and K. hoeninghausia from Germany rate morphotype or species. and K. cf. lamellosus and K. lamellosus from the Montagne Protoconch oval; diameter 0.68 mm (n = 13; range 0.58- Noire, France. Species of Koenenites matching the sequence 0.80 mm); width 0.63 mm (n = 10; range 0.58-0.70 mm). in the Penn Yan, Genundewa, and West River of New York Weak constriction between 0.75 and 1.0 volutions. Suture occur with equivalent zone conodonts in the Harrell Shale with broad, rounded primary lobes and saddles through two of Central Pennsylvania (Milesburg-Unionville, Center volutions. External or ventral lobe subdividing (E1-Em-E1) County) (Kirchgasser, 1996b). between 2 and 2.75 volutions (2.5-4.5 mm diameter) with Range.–Koenenites Genozone UD I-B to Timanites Em arising in flank of ventral lobe in “pharciceratid” style Genozone UD I-C. Conodont Zones MN 2-4. (Bensaïd, 1974) rather than “manticoceratid” style, in which E1 lobe arises from lengthening of Em saddle. Shallow inter- Koenenites styliophilus styliophilus (Clarke, 1898) nal U2 lobe seen at 8 mm diameter (3.5 volutions). Pl. 8, Figs 1-4, 6, 9; Text-figs 34A-F, I-J, L, 35A; Table 8 Diagnostic external umbilical lobe (U2) of Koenenites normally seen between 15 and 20 mm diameter (fourth and Manticoceras pattersoni var. styliophilum Clarke, 1898, 1899a, b: fifth volutions) but delayed or incipient in some specimens 47-48, 75, 82, text-fig. 55, pl. 6, fig. 30 (not text-fig. 2= K. (not seen at 20 mm diameter in NYSM 16571). Lateral sad- n. ssp.). styliophilus kilfoylei dle relatively large and asymmetrical, with steep dorsal side Manticoceras styliophilum Clarke. Miller, 1938: 119-121, fig. 24B (not fig. 24A = K. styliophilus kilfoylei n. ssp.), pl. 20, fig. 5. between 20 and 30 mm diameter, but thereafter becoming more symmetrical and distinctly flat-topped (Text-fig. 34F) Type specimen.–Lectotype designated herein is NYSM as lateral lobe becomes narrowly rounded to acute. 3783, the surviving syntype (Clarke, 1899: pl. 6, fig. 30; Growth lines and ornament distinctive. Swellings or ?text-fig. 55), from uncertain horizon and locality but prob- nodes developing on flanks near end of first volution, be- ably from the upper Penn Yan Shale in the region of Naples coming prominent in second volution, weakening through (Ontario County). third and becoming reduced to narrow ribs or ridges. Diag- New material.–Nearly 300 specimens from 14 localities nostic raised lirae of the genus Koenenites arising from ridge in the upper Penn Yan Shale and equivalents in the middle crests and asymmetrical with steep aboral faces. Bands of Ithaca Shale and Sandstone. The majority are fragments of finer, normal growth lines between lirae. Pattern of raised molds or calcitic replacements; some are pyritic or baritic lirae and growth lines showing broad salient centered low replacements. Included are 17 specimens in the NYSM from on flanks and moderately deep saddle centered dorsad of the Crosby Sandstone at Keuka Lake collected by D. Luther mid-flank. Strong, narrowly curved salient of raised lirae at in 1904 (his locality 3397). Mentioned and figured material ventrolateral margin, seen as crescent-shaped ribs in internal includes NYSM 3783, 12134-12142, 12158-12161, and molds (Pl. 8, Fig. 3). Growth lines and lirae crossing venter 16571. in high, rounded saddle. Dimensions.–See Table 8. Dorsal wrinkle layer, seen in baritized specimens, con- Description.–Moderately evolute; UW/D between 0.4 sisting of closely spaced, wavy lines in pattern similar to that and 0.3 mm to approximately 6 mm diameter, decreasing to in Manticoceras (House, 1971: pl. 1, figs 1-2). Ventral wrin- approximately 0.24 at higher diameters (Text-fig. 35A; Table kle layer consisting of closely spaced, continuous, parallel, 8). Whorl form subcircular (WW/WH=1) to 7-8 mm di- but moderately wavy lirae that slant gently backward across ameter (3-4 volutions) becoming broadly subtriangular be- umbilical wall becoming rectiradiate across flanks and ven- tween 10 and 20 mm diameter. Maximum whorl width ter. (WW) at top of steep umbilical wall with flanks converging Discussion.–Clarke regarded Manticoceras styliophilum as to broadly rounded venter (Text-figs 34A-E). Beyond 20 the “. . . characteristic variation of the specific type [Mantic- mm diameter, umbilical wall becoming less steep, flanks oceras pattersoni = M. sinuosum] most abundantly developed more flat-sided, and venter subtabular (Text-figs 34A-B). in the Styliola Limestone [Genundewa Limestone] . . . most Largest specimen observed (NYSM 12140) approximately from . . . outcrops . . . on Canandaigua Lake, and a single 55 mm diameter with body-whorl length of at least one-half representative . . . in the Naples fauna.” (Clarke, 1899a, b: volution. 75). Clarke only figured two specimens, an uncataloged and A few specimens appear to be considerably more de- now lost specimen from the Genundewa Limestone [Clarke, pressed and broader in whorl form than typical forms (Text- 1899a, b: 47, text-fig. 2; refigured herein (Text-fig. 34G) as 114 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

Text-fig. 33. Sutures and cross sections of Koenenitidae, Acanthoclymeniidae and Gephuroceratidae from the middle and upper Genesee Group of New York. A-E. Koenenites beckeri n. sp. from the West River Shale. A-C. Specimens from a loose concretion near the Bluff Point Siltstone (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 specimen from a concretionary level 0.67 m (2.2 ft) below the Bluff Point (Loc. 23a/3), Seneca Point Creek, Canandaigua Lake, (continued right) HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 115

Koenenites styliophilus kilfoylei n. ssp.], and the syntype from 4) and labeled as from the Genundewa Limestone at the Naples fauna (NYSM 3783) (Clarke, 1899a, b: pl. 6, Taunton Gully in the Genesee Valley, but that almost cer- fig. 30, text-fig. 55; refigured herein as Pl. 8, Figs 1-2) that tainly came from the Linden Horizon (Loc. 17/6) or there- is the designated lectotype. The horizon and location of the abouts in the upper Penn Yan Shale (J. W. Wells, pers. specimen is uncertain. Written in script on the original label comm., 1974). attached to NYSM 3783 are the words “Gon. styliophilus NYSM 3783 also closely matches specimens of compa- Naples concr.” indicating that the specimen came from a rable size from the Crosby Sandstone of the upper Penn Yan concretion or concretion horizon but it is not clear whether Shale in the vinicity of Keuka Lake, including a collection “Naples” refers to the Naples fauna, a designation that reported as Manticoceras from Sartwell Ravine (Loc. C, Py- would include horizons within the Genesee Formation, the 11) by D. D. Luther in 1904. Luther’s loc. 3397 at 880 ft Naples Group or Beds or Formation of older terminology (268 m) elevation, 1 mi south of Penn Yan, miscorrelated as that lie above the Genesee in the Sonyea Group, or the vil- Cashaqua Shale, is the Crosby Sandstone (Kirchgasser, lage of Naples in Ontario County. Two labels with the spec- 1985). Several specimens among the 17 in the NYSM col- imen indicate the locality as Naples beds, Naples, and in the lection have the same distinctive shell form and growth-line NYSM catalog the specimen is listed as from the “Portage pattern of NYSM 3783, and one in particular of similar di- (Naples) beds, Naples, New York.” It is important to note mensions (NYSM 16563, Pl. 8, Fig. 9) shows the suture that the limits of the Styliola or Genundewa Limestone were with the unmistakable U2 umbilical lobe of Koenenites. never precisely defined by Clarke in its type area at The lost syntype figured by Clarke (1899a, b: 47, text- Canandaigua Lake, and he probably included in the Styliola fig. 2; refigured herein as Text-fig. 34G) probably came from Limestone the lithologically similar styliolinid limestones of the Genundewa Limestone at Canandaigua Lake and is here the upper Penn Yan Shale. Furthermore Clarke’s Naples included as Koenenites styliophilus kilfoylei n. ssp. fauna included the fauna of the Genesee Group. We believe Rare specimens with a distinctly different sutural pattern that the surviving syntype of M. pattersoni var. styliophilum from that described above have been noted in the Crosby (NYSM 3783), herein determined as K. styliophilus stylio- Sandstone and are referred to morphotype B [e.g., NYSM philus, came from the Penn Yan Shale in the region of 16572-16574 (Loc. F, Py-16) and NYSM 12141 (Text-fig. Canandaigua Lake. Clarke’s Genundewa representatives are 34J)]. In morphotype B specimens, the sutures are closer to- the related form, K. styliophilus kilfoylei n. ssp. gether and in the adult the lateral saddle is asymmetrical and The preservation of NYSM 3783, with a thick shell wall not flat-topped as in the more common morphotype A. Also of recrystallized black calcite and raised growth-line lirae, is the lateral lobe of morphotype B is more angular and lies similar to the specimens described herein from styliolinid more ventrad of the less-steep umbilical wall. The stage of beds in the upper Penn Yan Shale, and especially from the appearance of the U2 lobe in morphotype B also varies. In Linden Horizon, the Crosby Sandstone, and equivalent NYSM 16573 it is seen at approximately 27 mm diameter horizons in the Ithaca Shale and Sandstone (Kirchgasser, but in NYSM 16574 is not seen at 35 mm diameter. Mor- 1985). NYSM 3783 is remarkably similar to a specimen col- photypes A and B do not appear to differ in other features lected by J. W. Wells (CU 40101, figured herein, Pl. 8, Fig. and 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 specimen from loose blocks in an embankment below NY Rte. 20 (Loc. 24/1) at Bethany Center, Genesee County. G. NYSM 12146, cross section at 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 the West River Shale at Hicks Point (Loc. 29a), Canandaigua Lake, Ontario County; figured by Wells (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 horizon in 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, 2.4 m (8 ft) below the base of the Middlesex Shale, in Beards Creek, Leicester, Livingston County, X 9.6. N. K. styliophilus kilfoylei n. ssp. from the Genundewa Limestone, NYSM 12147, whorl form and sutures (reversed) of a specimen from Loc.24/1, X 2.2. O. K. styliophilus kilfoylei n. 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. 116 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

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), recorded as 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) immediately below the Linden Horizon in Taunton Gully, Livingston County. C. NYSM 12137, cross section at 16 mm diameter of a specimen from the Linden Horizon (Loc. 15/8), Penn Yan Shale, Linden, Genesee County. D. NYSM 12138, cross section at 10.1 mm of a specimen from the Linden Horizon (Loc. 15/8), at the same locality. E. K. styliophylus styliophilus?, NYSM 12142, cross section at ca. 33 mm diameter of a specimen from the Penn Yan Shale (Loc. 17a/3), Spezzano Gully, Livingston County. F. K. styliophilus styliophilus, NYSM 12139, suture at 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) HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 117

Table 9. Biometric data for Koenenites cooperi Miller, 1938, from the Squaw Bay Limestone near Alpena, Michigan.

Specimen D WW WH UW WW/WH UW/D

MMP 13846 ca. 72.0 ca. 17.0 31.8 ca. 18.0 0.53 0.25 ca. 58.0 14.5 — 13.5 — 0.23 41.1 — 21.5 —— — 38.0 — 17.3 8.0 — 0.21 35.4 ca. 10.8 17.8 — 0.61 — 32.1 — 16.1 7.5 — 0.23 31.5 — 14.3 7.8 — 0.25 ca. 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 0.24 24.0 7.7 8.0 5.4 0.96 0.23 19.4 6.4 9.1 4.2 0.70 0.22 14.6 5.2 6.6 3.6 0.79 0.25 USNM 96546c 38.4 11.5 19.1 7.4 0.60 0.19 USNM 96546d ca. 68.0 ca. 18.0 29.0 17.0 0.62 0.25 USNM 96546f 16.0 6.0 6.8 4.6 0.88 0.29

rate species. In suture pattern, morphotype B resembles Koe- angular. In these features, K. lamellosus more closely resem- nenites styliophilus kilfoylei n. ssp. from the lower Genundewa bles K. styliophilus kilfoylei n. ssp. from the Genundewa Limestone to which it might be allied. Limestone and especially K. beckeri n. sp. from the West Koenenites styliophilus styliophilus is similar to K. cooperi River Shale. from the Squaw Bay Limestone of Michigan. In K. cooperi Koenenites lamellosus kirchgasseri from the Harrell Shale (see biometric data in Table 9), the adult whorl form is wider near Landes (Grant County), West Virginia, has a much and more broadly rounded across the venter. The sutural more lingulate lateral saddle and U2 lobe than K. styliophilus ontogenies are closely comparable but in the adult suture of styliophilus and in K. sp. of House (1978), also from the Har- K. cooperi, the lateral saddle becomes high and narrow and rell Shales near Landes. The whorl form is broader and the strongly asymmetrical (Miller, 1938: fig. 26B); at compara- external sutural elements are more narrowly rounded than in ble stages in K. styliophilus styliophilus, the lateral saddle is K. styliophilus styliophilus. These West Virginia forms pre- broader, more symmetrical, and distinctly flat-topped. The sumably represent an earlier stage in the evolution of Koe- suture of K. cooperi is somewhat closer in form to morpho- nenites than the Penn Yan and Genundewa representatives type B and its suture and whorl form most closely resemble from New York. K. styliophilus kilfoylei n. ssp. from the Genundewa Lime- Distribution.–Upper Penn Yan Shale and equivalents in stone. Ithaca Shale and Sandstone (Genesee Group; Wyoming to Compared to Koenenites styliophilus styliophilus the types Tompkins counties). of K. lamellosus from Oberscheld, Germany, have a rather Koenenites styliophilus styliophilus is restricted to the broader whorl form across the flanks and venter and a suture upper Penn Yan Shale and is best represented in the Linden with a higher, narrower, and U-shaped lateral saddle flanked Horizon and surrounding styliolinid beds, the equivalent or by deeper E1 and U2 lobes with the latter being distinctly slightly 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 Canandaigua Lake. Enlarged but scale unknown. H. K. beckeri n. sp., NYSM 12156, partial whorl form of a specimen from a loose concretion near the Bluff Point Siltstone (Loc. XX, Nap-8/1) of the West 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 mm diameter 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 Beards Creek Horizon (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. L. Same as F. Scales = 5 mm. 118 B LEISO MRCNPLOTLG o 374 no. PALEONTOLOGY AMERICAN OF ULLETINS

Text-fig. 35. Graphs showing the ontogeny of Koenenites, Manticoceras, and Probeloceras spp. A. Koenenites styliophilus styliophilus (Clarke, 1898), based on specimens from the Penn Yan Shale; based on data in Table 8. B. Manticoceras contractum (Clarke, 1898), based on a cotype (NYSM 5125) from the Genundewa Limestone; M. nodifer (Clarke, 1885), based on type material, NYSM 14994a,b, from the Genundewa Limestone; and M. sinuosum apprimatum Clarke, 1898, based on type material (NYSM 3732) from the Genundewa Lime- stone. C. Probeloceras lutheri (Clarke, 1885), based on data from material from the Cashaqua Shale in Table 22. HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 119

Table 10. Biometric data for Koenenites styliophilus kilfoylei n. ssp. from the Genundewa Limestone.

Specimen D WW WH UW WW/WH UW/D

NYSM 12162 34.5 ca. 10 15 8.5 0.67 0.25 23.3 ca. 8 10.5 6 0.76 0.26 NYSM 12144 ca. 27 — 12.5 ca. 8.5 — 0.31 NYSM 12145 26.7 9 12 6.8 0.75 0.25 18 ca. 6.5 8.3 4.2 ca. 0.78 0.23 11.6 ca. 4.8 5.6 2.7 ca. 0.86 0.23 NYSM 12146 26.0 ca. 10 11.3 7 0.88 0.27 17.2 ca. 7.3 7.4 4.5 0.99 0.26 11.6 5.4 5.1 3.3 1.06 0.28 7.8 3.4 3.3 2.5 1.03 0.32 5.2 2.5 2.0 2.1 1.25 0.40 3.8 1.6 1.3 1.7 1.23 0.45 NYSM 12163 26 9.2 11.8 6.6 0.77 0.25 17 6.4 7.6 5.3 0.84 0.31 NYSM 12164 19 ca. 6.5 8.4 5.3 0.77 0.27 ca. 13.3 — 5.7 4.3 — 0.32

sentatives or equivalents of these horizons in the Ithaca For- Loc. EE, Py-14); Sunset Point (Loc. FF, Py-16), creek above mation at Seneca and Cayuga lakes (Seneca and Tompkins Sunset Point (Yates County). Mentioned or figured material counties; Kirchgasser, 1985). includes NYSM 12143-12147, and 12162-12165. Range.–Frasnian: Koenenites Genozone UD I-B. Re- Dimensions.–See Table 10. gional Zone of K. styliophilus styliophilus (16a). Conodont Diagnosis.–Closely similar to nominate form but slightly Zone MN 2. more evolute, broader in whorl form in early stage, more flat-sided on flanks, with more broadly rounded venter and Koenenites styliophilus kilfoylei n. ssp. suture with asymmetrical lateral saddle that is not flat- Pl. 8, Figs 5, 7-8, 12-13; Text-figs 33F, G, N-O, 34G; Table 10 topped. Description.–Shell form and suture similar to nominate Manticoceras pattersoni var. styliophilum Clarke, 1899: 47, text-fig. forms from Penn Yan Shale but slightly more evolute and 2. broader in cross section in early stages (WW/WH higher) aff. House & Kirchgasser, 1993: 276-277. Koenenites styliophilus and becoming more flat-sided on flanks and more broadly rounded across venter (Text-figs 33F-G, N, 34G). Suture Etymology.–Named in honor of Clinton Kilfoyle, former similar to nominate form in having broadly rounded lateral Curator of Paleontology at the New York State Museum. saddle. Lateral saddle compares most closely with morpho- Type specimen.–NYSM 12143 (Pl. 8, Figs 7-8), from the type B of Koenenites styliophilus styliophilus in having lateral lower 15 cm of the Genundewa Limestone, from loose saddle more asymmetrical and not as flat-topped as mor- blocks at NY Rte. 20 embankment at Bethany Center photype A of K. styliophilus styliophilus (Text-figs.33N, 34J). (Genesee County; Loc. 24/1). The lost syntype and uncat- Included here is single morphotype (form A, NYSM 12144) aloged specimen of Manticoceras pattersoni var. styliophilum with unusual suture of closely spaced, tongue-shaped lateral from the Genundewa Limestone figured by Clarke (1899a, saddles with rounded crests (Pl. 8, Figs 12-13); lateral saddle b: text-fig. 2; refigured herein, Text-fig. 34G) is included in narrower and more symmetrical than typical K. styliophilus Koenenites styliophilus kilfoylei n. ssp. kilfoylei n. ssp. in lower Genundewa and not as flat-topped Material.–Over 100 specimens from the lower Genun- as typical form of K. styliophilus from upper Penn Yan Shale. dewa Limestone: 74 from loose blocks of the lower 15 cm As in Koenenites styliophilus styliophilus, stage of appear- of the unit at the NY Rte. 20 embankment, Bethany Center ance of U lobe varies. In NYSM 12147, seen at approxi- (Genesee County; Loc. 24/1); 6 from Murder Creek, Gris- 2 mately 18 mm diameter (Text-fig. 33N) and in NYSM wold (Genesee County; Loc. 14a/3); 23 from Linden Falls, 12144, at approximately 27 mm (Text-fig. 33O), but not Linden (Genesee County; Loc. 15a/12); 4 from Beards in NYSM 12165 at approximately 26 mm diameter. Creek, Leicester (Livingston County; Loc. 18a/6a); several Growth lines and wrinkle layer similar to K. styliophilus from the pyritic nodular horizon equivalent to the Genun- styliophilus but raised growth-line lirae not as strongly de- dewa near Keuka Lake: Cornwall Gully, Penn Yan (Yates veloped. County; Loc. DD, Py-10); Willow Grove (Yates County; 120 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

Discussion.–Koenenites styliophilus kilfoylei n. ssp. com- show moderately evolute coil with deep umbilicus within pares closely with morphotype B of K. styliophilus styliophilus which distinctive, paired, fasciculate growth lines are dis- of the upper Penn Yan Shale and is a likely descendant of played. These features clearly seen in NYSM 16575 from that taxon. Close similarity with K. cooperi of the Squaw Bay Genundewa Limestone at Genundewa Point (Loc. 23b/4) Limestone, Michigan, is seen in whorl form and suture but and in baritic specimen (NYSM 16590) from Genundewa in K. cooperi the asymmetry of the lateral saddle is stronger Pyrite at Sunset Point (Loc. FF, Py-16/2). and the external U2 lobe more fully developed. Similarly the Innermost whorls, preserved as pyritic replacements in suture of K. lamellosus is more advanced in having higher types and new material from Genundewa Pyrite, have lateral saddles and lobes and more strongly developed U2 strongly depressed whorl form with broadly rounded venter lobe. and distinctive, broad, rib-like swellings on flanks (Miller, Distribution.–Lower Genundewa Limestone (Genesee 1938: fig. 15). Swellings appear to mark sites of raised, fas- Group) in Genesee and Livingston counties and equivalent ciculate growth lines seen in calcitic and baritic replace- pyritic horizon near Keuka Lake in Yates County and in ments. equivalents of the Genundewa Limestone in the Harrell Discussion.–The immature suture is distinguished by a Shale at Milesburg and Unionville (Center County), Penn- large, prominent ventral lobe from the sides of which the sylvania. ventrolateral lobes develop in the fourth whorl. This pattern Range.–Frasnian: Koenenites Genozone UD I-B Regional of sutural development on the venter, seen in the syntypes Zone of K. styliophilus kilfoylei (16b). Conodont Zone MN and new material, was recognized as primitive by Clarke 2. (1898) and unlike the pattern seen in Manticoceras in which the ventrolateral lobes arise from the enlargement of the E1 Koenenites? fasciculatus (Clarke, 1898) saddles. This pharciceratid-type of evolving external lobe is Pl. 18, Figs 1-2, 5; Text-figs 33I-J also seen in Pseudoprobeloceras, Acanthoclymenia, other species of Koenenites, and probably also Hoeninghausia (Ben- Manticoceras fasciculatum Clarke, 1898, 1899a, b: 71-74, 81-82, saïd, 1974). Because of their small size, the syntypes of K.? pl. 6, figs 13-22, text-figs 45-53, 63-64. fasciculatus provide no information about the mature suture. Clarke. Miller, 1938: 85-87, text-fig. 15, Manticoceras fasciculatum One of the specimens (NYSM 12155), however, from the pl. 15, figs 1-10. Genundewa Pyrite at Cornwall Gully (Loc. Py-10/1) shows the suture of a Koenenites at approximately 51 mm diameter Types.–Nine syntypes of this species are housed in the with a deep lateral lobe, high saddle across the umbilical New York State Museum as NYSM 3736-3744. All are small wall, and a clearly defined U lobe outside the seam (Text- juveniles of less than 5 mm diameter. According to Clarke 2 fig. 33J). The pyritic inner whorls of the specimen are poorly (1898), all are from the Genundewa Limestone at preserved and although the irregular wrinkle-layer pattern Canandaigua Lake and Middlesex (Yates County). A lecto- of Koenenites is seen, the typical rib-like swellings of Clarke’s type is here designated as NYSM 3738 (Pl. 18, Figs 1-2), species Manticoceras fasciculatum are not seen. Thus the from the Genundewa at Canandaigua Lake. generic assignment of K.? fasciculatus and the assignment of New material.–Eleven specimens from the Genundewa NYSM 12155 to the species are problematic. Limestone and pyritic nodular equivalent of the Genundewa Distribution.–Lower Genundewa Limestone (Genesee (Genundewa Pyrite) near Keuka Lake (Yates County); one Group), Canandaigua Lake (Yates County). Also in Genun- calcitic replacement (NYSM 16575) of the inner whorls dewa Pyrite equivalent of the Genundewa near Keuka Lake from the Genundewa Limestone at Genundewa Point (Yates (Yates County). County; Loc. 23b/4), 0.71 m above the base of the unit; Range.–Frasnian: Koenenites Genozone UD I-B; ?Timan- two partly pyritic crushed molds of adult specimens from ites Genozone UD I-C. Regional Zone of K. styliophilus kil- the Genundewa Pyrite at Cornwall Gully, Penn Yan (Yates foylei (16b), ?Manticoceras contractum (17a). Conodont County; Loc. DD, Py-10); seven pyritic and one baritic re- Zone MN 2-?3. placements of inner whorls from the Genundewa Pyrite at Sunset Point Gully, Keuka Lake (Yates County; Loc. FF, Py- Koenenites? cf. fasciculatus (Clarke, 1898) 16). Description.–Like syntypes, most new specimens are Material.–Thirty-three pyritic inner whorls recovered small inner whorls and consequently little can be added to from a loose concretion, probably from the first rhythm in Clarke’s (1898) descriptions and illustrations summarized the West River Shale beneath the Bluff Point Siltstone in a by Miller (1938). Largest types [Clarke, 1899a, b: pl. 6, fig. gully 0.8 mi (1.3 km) south of Middlesex (Yates County; 19 (lost syntype) and 21 (NYSM 3738)] at four volutions HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 121

Loc. XX, Nap-8/1). One pyritic replacement of an inner shales beneath a black shale near the level of the Bluff Point whorl from the Fossil-Log concretion horizon in the upper Siltstone at 1,030 ft (314 m) elevation and 30 m below the West River Shale (Williamsburg Bed of Over et al., 1999, Middlesex in Sheldrake Creek, Cayuga Lake (Seneca 2003), 3.4 m below the Middlesex Shale in Beards Creek, County; Loc. MM, Gen-1a); one mold in a silty shale in the Leicester (Livingston County; Loc. 18a/4b). Ithaca Shale and Sandstone near the level of the Bluff Point Discussion.–The quality of preservation varies widely but Siltstone at approximately 980 ft (299 m) elevation and 47 all of the specimens conform closely in whorl form and su- m below the Middlesex in Lick Brook, Ithaca (Tompkins ture to Clarke’s types of Koenenites? fasciculatus and the new County; Loc. OO, I-8); two molds from a gray shale 4.9 m specimens assigned to that species from the lower Genun- below the Middlesex Shale in Beards Creek, Leicester (Liv- dewa Limestone and equivalent Genundewa Pyrite in the ingston County, Loc. 18a/4a); one calcite and barite replace- vinicity of Keuka Lake (Yates County). It is uncertain ment from a concretion horizon 5 m below the Middlesex whether these inner whorls from the middle and upper West in Snyder’s Gully, Woodville, Canandaigua Lake (Ontario River Shale are the same as K.? fasciculatus from the Genun- County; Loc. 46/c1). dewa Limestone or whether they are early stages of the large Diagnosis.–Closely similar to Koenenites lamellosus but form from the same horizons referred here to K. beckeri n. with more flat-sided to nearly parallel-sided flanks and sp. broader to nearly flat venter. Later saddle of K. beckeri n. sp. Distribution.–Middle and upper West River Shale is slightly more angular than in K. lamellosus. (Genesee Group) in Livingston and Yates counties. Description.–Shell form of large specimens moderately Range.–Frasnian: Timanites Genozone UD I-C. Re- evolute with flattened and nearly parallel flanks that slope gional Zone of Koenenites beckeri (17b). Conodont Zone gently to broadly rounded to flattened venter. Somewhat MN 4. rectangular cross section of flanks and venter in largest spec- imens is distinctive. Koenenites beckeri n. sp. Adult suture with high, strongly asymmetrical lateral Pl. 8, Figs 10-11, 14; Text-figs 33A-E, 34H, K saddle, narrowly rounded to angular lateral lobe, relatively high saddle on umbilical wall, and shallow U2 lobe on or Type material.–Holotype here designated, NYSM 12150 just outside umbilical seam. Growth lines with raised lirae (Pl. 8, Fig. 14; Text-fig. 33C) from the rhythm below the typical of Koenenites that, where seen on inner whorls, are Bluff Point Siltstone near Middlesex (Loc. XX, Nap-8 of de somewhat fasciculate. On flanks of largest specimens, Witt & Colton, 1978). growth lines show sinus on flank and strong tongue-like ven- Etymology.–Named in honor of our colleague, R. T. trolateral salient and broadly rounded sinus across venter. Becker, University of Münster, Germany. Discussion.–The whorl form and suture indicate an affin- Previously referred to Koenenites sp. C (Kirchgasser & ity with Koenenites lamellosus. In the types of K. lamellosus House, 1981: 42, text-fig. 2) and K. aff. lamellosus (see from Oberscheld, Germany, however, the flanks are not as House & Kirchgasser, 1993: 277, fig. 7). flat-sided and the lateral saddle is more rounded than in the Additional material.–Twelve specimens from the rhythm New York specimens at comparable stages. below the Bluff Point Siltstone of the middle West River The New York specimens are also similar to the speci- Shale and the correlative horizon in the Ithaca Shale and mens referred to Koenenites cf. lamellosus and K. lamellosus by Sandstone and two specimens from the upper West River House et al. (1985: figs 8A-C) from Bed 58 in the CPS-E Shale; three calcitic replacements/molds (including NYSM section in the Montagne Noire, France, but again they differ 12153) from the gray shale in the first rhythm beneath the in having flatter flanks at comparable stages. The specimen Bluff Point Siltstone (0.76 m below the horizon) at Seneca referred to Hoeninghausia cf. archiaci from the slightly Point Creek, Canandaigua Lake (Ontario County; Loc. younger Bed 24 in Trench A’ at La Serre, in the Montagne 23a/3); six calcitic replacements (including the holotype Noire, has a similar suture (House et al., 1985: pl. 3, figs NYSM 12150, and 12149, 12151-12153, 12156) from a 10-12, text-figs 8D-E) but at a stage twice the size of the loose concretion probably from the same first rhythm be- New York specimens the flanks slope much more steeply to neath the Bluff Point Siltstone in a gully 0.8 mi (1.3 km) a more narrowly rounded venter. As noted earlier, the genus south of Middlesex (Yates County; Loc. XX, Nap-8/1); two Hoeninghausia (type H. archiaci) is restricted to undoubtedly molds from a dark gray shale beneath the horizon of the oxyconic forms. The specimen from Bed 24 in Trench A’ at Bluff Point Siltstone at 920 ft (280 m) elevation and 24 m La Serre is a transitional form best referred to K. cf. lamel- below the Middlesex Shale in Mill Creek, near Lodi (Seneca losus. County; Loc. LL, OV-10/16); one mold from dark gray The type specimen of Ammonites hoeninghausia from 122 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

Bensberg, Germany, which was reillustrated and assigned to Family GEPHUROCERATIDAE Frech, 1897 Hoeninghausia (Koenenites) hoeninghausia by Bensaïd (1974: The gephuroceratids comprise the family named by 107, pl. 3, fig. 7, text-fig. 14), shows a remarkable similarity Wedekind (1913) as Manticoceratidae after its best known to the New York specimens of comparable size in its flat- genus, Manticoceras, but the earlier name of Frech has pri- tened flanks and broadly flattened venter and acute ventral ority and widest use. The group is thought to have arisen lobe, but its U2 lobe is acute as well. The horizon of von from late Givetian serpenticonic acanthoclymeniids such as Buch’s specimen is unknown but its suture suggests a more Pseudoprobeloceras by a tendency toward a more involute advanced member of the K. lamellosus lineage than the type shell with very varied morphology, an emphasis of the mid- of the species and the specimens described herein as K. beck- dorsal saddle, and the addition of umbilical lobes. Growth eri n. sp. from the middle and upper West River Shale. lines are biconvex but in some later members (Crickites) be- Distribution.–Middle and upper West River Shale come convex. The family characterizes the Frasnian world- (Genesee Group) in Livingston, Ontario, Yates, Seneca, and wide but became extinct at its close with the Upper Tompkins counties. Kellwasser Event (Becker & House, 1993, 1994a). Range.–Frasnian: Timanites Genozone UD I-C. Re- The family has three named subfamilies, Gephurocerati- gional Zone of Koenenites beckeri (17b). Conodont Zone nae, Virginoceratinae, and Crickitinae, but the characters of MN 4. the first group are so varied as to indicate the division is rather an arbitrary one. Koenenites beckeri n. sp., form D Included genera in the Gephuroceratinae are Mantico- Text-figs 33E, 34K ceras Hyatt, 1884, Clauseniceras Becker & House, 1993, Costamanticoceras Becker & House, 1993, Delphiceras Discussion.–We previously referred to Koenenites sp. D Becker & House, 2000 (replacement for Delphinites Becker (Kirchgasser & House, 1981: 43, text-fig. 2) and K. aff. & House, 1993, non Sayn, 1901), Maternoceras Clausen, lamellosus sp. D (House & Kirchgasser, 1993: 277, text-fig. 1971, Playfordites Becker et al., 1993, Trimanticoceras House 7), two specimens (NYSM 12154 and 12157) from the in House & Ziegler, 1977, Mixomanticoceras Becker et al., Beards Creek Horizon (Bed 18a/5) in upper West River 1993, Sphaeromanticoceras Clausen, 1971, and Timanoceras Shale, 2.4 m below the Middlesex Shale, at Beards Creek, Bogoslovsky, 1957. The four known in New York are Man- Leicester (Livingston County; Loc. 18a). NYSM 12157 ticoceras, Delphiceras, Playfordites, and Sphaeromanticoceras. (Text-fig. 34K), which we regard as the type, is a fragment Included in the subfamily Virginoceratinae Yatskov, of the outer whorl of a large individual. The raised growth- 1990, are the genera Virginoceras Ruzhencev, 1960, Carino- line lirae seen on the outer flank is of the Koenenites type. In ceras Lyashenko, 1957, and Neomanticoceras Schindewolf, cross section, the flanks slope to a broadly rounded venter 1936. Of these only Carinoceras is recognized in New York. giving a distinctly different profile than the parallel-sided The subfamily Crickitinae Wedekind, 1913, is charac- profile with broadly flattened venter seen in the specimens terized by convex growth lines from an early stage. Crickites lower down in the West River Shale referred to K. beckeri n. occurs in New York and Canada. sp. Although the specimen is crushed, the cross section of In the text below, genera are treated in their approximate NYSM 12154 also shows flanks that are not subparallel but order of appearance in New York. clearly slope toward the venter. Distribution.–Worldwide. Assigned to the same taxon is a third specimen (NYSM Range.–Frasnian: Manticoceras Stufe: ?Koenenites Geno- 16576) that shows a similar profile but a somewhat more zone UD I-B to Crickites Genozone UD I-L. Conodont broadly flattened venter. Specimen NYSM 16576 is from Zones MN 3-13. the fossil log concretion horizon in the upper West River Shale, 3.4 m below the Middlesex Shale in Chidsey Point Subfamily GEPHUROCERATINAE Frech, 1897 Creek, Keuka Lake (Steuben County; Loc. KK, Py-7/2; Genus MANTICOCERAS Hyatt, 1884 Williamsburgh Bed of Over et al., 1999, 2003). Type species.–Goniatites simulator Hall (1874: 2), by Distribution.–Upper West River Shale (Genesee Group) monotypy. in Livingston and Steuben counties. Diagnosis.–Conch usually large, evolute in inner whorls Range.–Frasnian: Timanites Genozone UD I-C. Re- then subevolute to subinvolute. Shell without ribs or nodes; gional Zone of Koenenites beckeri (17b). Conodont Zone growth lines biconvex. Suture with three pointed or rounded MN 4. lobes between small ventral lobe (Em), and umbilical seam: saddle between E1 and L (Text-fig. 25) always large and high. HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 123

Included species.–Even after the recent separation of sev- characterize this species. Specimen 3872/9 from the Genun- eral genera, there are still over 40 specific names that could dewa at Fall Brook (Loc. 19a) has a growth-line pattern and be used in this genus. Only those relevant to the description broadly rounded venter that matches the features of the des- of New York taxa will be referred to below. A full review of ignated lectotype (NYSM 3735). NYSM 5125 at 22.3 mm the genus is needed. has a rather narrow whorl-form (Miller 1938: pl.13, fig. 14) Discussion.–Aspects of the taxonomy of the genus were and it could be a Koenenites from the upper Penn Yan Shale. reviewed by Miller (1938: 72) and the evolutionary relations Distribution.–Upper Genundewa Limestone (Genesee of the group have been discussed by Becker & House Group) in Livingston and Yates counties. (1993). We follow Miller in considering Gephuroceras Hyatt, Range.–Frasnian: Timanites Genozone UD-I-C. Re- 1884, and Gephyroceras as junior synonyms of Manticoceras. gional Zone of Manticoceras contractum (17a). Conodont Distribution.–Worldwide except for polar regions and Zone MN 3. South America. New York: Upper Genundewa Limestone (Genesee Group) to Angola Shale (West Falls Group). Man- Manticoceras sinuosum apprimatum Clarke, 1898 ticoceras first appears in New York in the Upper Genundewa Pl. 17, Figs 5-10; Text-figs 35B, 42F Limestone with the entry of M. contractum, M. nodifer, and M. sinuosum apprimatum, the latter two also presumably Manticoceras apprimatum Clarke, 1898, 1899a, b: 62-63, 80, text- from the Upper Genundewa. figs 63-64, pl. 6, figs 27-29; Miller, 1938: 76, pl. 13, figs 9-11; Becker & House, 1993: 119. Range.–Timanites Genozone UD I-C to Crickites Geno- zone UD I-L. Conodont Zones MN 3-13. New York: Type material.–The three syntypes of Clarke (NYSM Timanites Genozone UD I-C to Neomanticoceras Genozone 3731-3733; formerly 12301/1-3) are refigured here and UD I-J. Conodont Zones MN 3-?11. NYSM 3732 is here designated as lectotype. Manticoceras contractum Clarke, 1898 Discussion.–Clarke’s types are poorly preserved; thus the Pl. 19, Figs 10-16; Text-fig. 35B characteristics of M. apprimatum will always be poorly un- derstood. The narrow umbilicus, broadly rounded whorl- Manticoceras contractum Clarke, 1898, 1899a, b: 69-70, 81, text- form, and gently sinuous biconvex growth lines seen in figs 44, 63-64, pl. 6, figs 1-2. NYSM 3732 suggest assignment to Manticoceras but unfor- Manticoceras contractum. Miller, 1938: 81-82, pl. 13, figs 12-14. tunately the suture is not seen in any of the types. It could well be the earliest representative of the M. sinuosum group Type material.–Three cotypes of Clarke (NYSM 3734, and tentatively we refer to it as a subspecies of that form. As 3735, and 5125) from the Genundewa Limestone near noted by Miller (1938), Clarke stated that this species is “not Middlesex (Yates County) and Canandaigua Lake (Yates of frequent occurrence.” NYSM 3731 (Pl. 17, Fig. 5), a frag- County). NYSM 3735 (Clarke, 1898: pl. 6, fig. 2; and ment showing its ribbed inner whorls, might be a Koenen- herein Pl. 19, Figs 12-15), Genundewa Limestone, ites. Canandaigua Lake, is here designated as lectotype. Distribution.–Genundewa Limestone (Genesee Group) New material.–Several specimens (some baritic) from in Yates and Genesee counties. The lectotype (NYSM 3732) loose blocks of the Genundewa at the base of the high falls and NYSM 3733 are recorded as from the Genundewa in Fall Brook (Loc. 19a), near Geneseo (Livingston County), Limestone at Middlesex (Yates County), and NYSM 3731, are assigned to this species (WK 3011, 3872); the blocks are the possible Koenenites, is from the Portage (Naples) Beds at believed to have come from the top of the Genundewa Griswold (Genesee County), which could be the Genun- Limestone. Three fragments (WK 3946/1/2/7) from Bed dewa Limestone in Murder Creek (Loc. 14a) at Griswold. 19/10, at the top of the nodular Lower Genundewa at Fall Range.–Frasnian: ?Timanites Genozone UD I-?C. ?Re- Brook-Dewey Hill (Loc. 19) are assigned with question to gional Zone of Manticoceras contratum (17a). Conodont this species. Zone uncertain, possibly MN 3. Diagnosis.–Summarized from Miller (1938) after Clarke (1898, 1899a, b): Small, subglobular conch with narrow Manticoceras nodifer (Clarke, 1885) umbilicus, rounded umbilical shoulders and ventrolateral Pl. 18, Figs 11-12; Text-fig. 35B margin, and broadly rounded venter. Ornamentation pecu- Goniatites nodifer Clarke, 1885: 21. liar with growth lines weakly biconvex and grouped into fas- Manticoceras nodifer. Clarke, 1898, 1899a, b: 74-75, 82, text-figs cicles or bundles at maturity (Pl. 19, Fig. 14). Suture 54, 63-64, pl. 6, figs 24-26; Miller, 1938: 92-93, pl. 15, figs unknown. 15-18; House, 1962: text-fig. 3G; Becker & House, 1993: 119, pl. 2, figs 2-3. Remarks.–Additional material will be required to better 124 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

Type material.–Clarke’s syntypes (USNM 14994a and 69, figs 1-2, pl. 74, fig. 8). Stated by Clarke (1899a, b) to be 1499b); lectotype selected by Becker & House (1993). the only specimen known to Hall; this is the only specimen Diagnosis.–Laterally compressed, subevolute shell with known to us. rounded venter, well-rounded umbilical shoulder, and steep Dimensions.–Maximum D = 42 mm; D = 37.2 mm; umbilical wall. Nodes on inner whorls numbering from 12 WW = 14.3 mm; WH = 17.2 mm; UW = 7.9 mm. per volution on inner whorls to 16 on third volution. Diagnosis.–Whorls compressed, very broadly rounded Growth lines not observed. Wrinkle layer in lectotype latterally, narrowly rounded ventrally, deeply impressed dor- consists of fine irregular striae numbering approximately 13 sally. Umbilicus deep and moderate in size, less than one- per mm on outer flanks at 10 mm diameter, which pass third diameter of shell. Mature suture with rounded lateral backward from umbilicus and sweep gently forward. lobes (Pl. 9, Fig. 5) (House, 1962: fig. 3F). Growth lines not Suture of lecotype (USNM 14994b) at 12 mm diameter, preserved. with U-shaped ventrolateral lobe, large asymmetrical lateral Description.–See Miller (1938: 105) for full description. saddle with orad portion of dorsal slope deepest, and broadly Discussion.–As noted by Miller (1938), the rounded lat- rounded lateral lobe (House, 1962: fig. 3G). eral lobes of the mature suture are distinctive. In typical Discussion.–There is some ambiguity as to which speci- species assigned to Manticoceras, the lateral lobes are narrow men was figured by Clarke (1898, 1899a, b: pl. 6, fig. 26) to pointed at comparable diameters. to show the inner whorls of this species; both specimens are Distribution.–Ithaca Shale and Sandstone (Genesee preserved as internal molds, partly of white calcite, in gray, Group), at or near Ithaca (Tompkins County). Precise hori- buff-weathered styliolinid limestone. The drawing appears zon unknown but thought to be from a level above the to be largely based on the lectotype but it is by no means an equivalent of the Lower Genundewa Limestone (Upper Ge- accurate copy because the artist gives the false impression nundewa and West River Shale). that the specimen is perfectly preserved. NYSM 3745 is a Range.–Frasnian: ?Timanites Genozone UD I-?C. ?Re- cast of the lectotype (USNM 14994b) and NYSM 11466 gional Zone of Manticoceras contractum (17a) or Koenenites (12304/1) is a cast of USNM 14994a. beckeri (17b). Distribution.–Upper Genundewa Limestone (Genesee Group), Genundewa Point, Canandaigua Lake (Yates Manticoceras sinuosum sinuosum (Hall, 1843) County; Loc. 23b; Clarke, 1885: 21). The horizon and col- Pl. 9, Figs 6-8, Pl. 10, Figs 1-12, Pl. 11, Figs 1-13, Pl. lection date of Clarke’s types are uncertain. The labels with 12, Figs 8-10, Pl. 15, Figs 1-2, Pl. 16, Figs 11-12; Text- the specimens indicate their collection by the Geological figs 36-38; Table 11. Survey in 1886 from the “Senecan (Genesee) of Lake Hall, 1843: 244, 246, text-figs 106(6), 107(9); Canandaigua.” More recent labels give the horizon as Ge- Goniatites sinuosus 1879 (pars): 460-463, pl. 72, fig. 11 (not pl. 70, figs 13, ?14, nundewa. In his original description, Clarke (1885: 21) gave pl. 74, fig. 11). the explicit location as “Genundewah [original spelling of Gon. Patersoni (sic) Hall, 1860: 99, text-figs 9-10. Genundewa], Canandaigua Lake.” He described it as a rare Manticoceras Pattersoni Clarke, 1898, 1899a, b (pars): 45-62, pl. 1, species and regretted that its “recent discovery” prevented figs 1-12, pl. 2, figs 1-2, 4 (not figs 3 = M. s. tardum, 6 = him from preparing “an illustration of it” in his 1885 report. Sphaeromanticoceras oxy), pl. 4, figs 16-17, ?18 (not figs 14-15 The horizon within the Genundewa Limestone at Genun- = Probeloceras lutheri?), text-figs 1, 3-11, ?12, 13-28. dewa Point (Loc. 23b) is unknown but a position in the Manticoceras sinuosum. Miller, 1938 (pars): 106-115, text-figs 1, 3- upper part of the unit is probable. 11, ?12, 13-28, pl. 18, fig. 4, pl. 19, figs 2-9 (not fig. 1 = M. Range.–Frasnian: ?Timanites Genozone UD I-?C. ?Re- s. tardum), pl. 20, figs 7-8, ?9, 10 (not fig. 6 = Probeloceras lutheri?); House, 1962: 259. gional Zone of Manticoceras contractum (17a). Type material.–The lectotype of Manticoceras sinuosum Manticoceras simulator (Hall, 1874) sinuosum is Goniatites sinuosus Hall [1843: 243, fig. 106(6) Pl. 9, Figs 4-5 (not 8) = AMNH 5887/1], selected by House (1962: 259), Goniatites simulator Hall, 1974: 2-3; Hall, 1875: 133-134; 1876: from the Cashaqua Shale, Cashaqua Creek (Livingston pl. 69, figs 1, 3, pl. 74, fig. 8; 1879: 453-455, pl. 69, figs 1-2, County). Among Clarke’s collection of Manticoceras Patter- pl. 74, fig. 8. soni are two fragments from the Naples Beds that have rather Manticoceras simulator. Hyatt, 1884: 318; House, 1962: text-fig. strong biconvex growth lines with a pronounced lateral 3F, pl. 45, fig. 14. sinus, narrow projecting ventrolateral salient and deep Type material.–NYSM 3797 (formerly 12309/1), men- ventral sinus (hyponomic recurvature): NYSM 3769 tioned by Hall (1874, 1875) and figured by him (1876: pl. (Clarke, 1899a, 1899b: pl. 4, fig. 14), Rock Stream (Yates HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 125

Table 11. Biometric data for Manticoceras sinuosum sinuosum (Hall, 1843) from the Cashaqua Shale. See Text-fig. 38 for graphs.

Specimen D WW WH UW WW/WH UW/D

NYSM 13835 62.5 20 33.5 9 0.60 0.14 NYSM 13800 58.5 ca. 22 ca. 30 ca. 9.5 0.73 0.16 36 14.2 19 6.4 0.75 0.18 22 9.5 10.9 4.6 0.87 0.21 13.9 6.6 6.5 3.6 1.01 0.26 8.8 4.4 3.7 2.9 1.19 0.33 6.0 3.0 2.1 2.3 1.43 0.38 4.0 2.0 1.4 1.8 1.43 0.45 2.9 1.4 0.90 1.4 1.56 0.48 2.1 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 0.21 ca. 11.3 ca. 5.5 ca. 5.3 3.1 1.04 0.27 7.0 ca. 3.5 3.1 2.4 1.13 0.34 4.6 ca. 2.4 1.7 2.1 1.41 0.46 3.3 1.5 1.1 1.2 1.36 0.36 2.4 ca. 1.1 0.75 1.1 1.47 0.46 NYSM 13811 44.4 16 ca. 23 8.0 0.70 0.18 NYSM 13801 38 14.7 20 6.4 0.74 0.17 ca. 23 10.2 ca. 11.1 ca. 5.0 0.92 0.22 14.6 6.9 7.0 3.8 0.99 0.26 9.2 4.7 3.9 3.0 1.21 0.33 6.1 3.3 2.3 2.5 1.43 0.41 4.2 2.2 1.4 1.9 1.57 0.45 3.1 1.6 1.0 1.4 1.60 0.45 2.3 1.1 0.74 1.0 1.49 0.43

County), and NYSM 3770 (Clarke, 1899a, 1899b: pl. 4, growth lines weakly sinuous with slight umbilical salient and fig. 15), Naples (Ontario County). This pattern of growth lateral sinus on mid-flank and distinctive, large, broad ven- line is not seen in M. s. sinuosum from the Cashaqua at any trolateral salient. stage and is more closely comparable with that seen in the Description.–Beginning at approximately 5.0 mm diam- largest specimens of Probeloceras lutheri. eter, whorl compression and impressment of dorsal area in- The spelling of pattersoni with two “t”s started with creasing abruptly; WW/WH ratio rapidly decreasing and Clarke (1898: 59) when he reported Hall as saying it was UW rapidly increasing (Text-figs 36-38). Typical chordate intended to honor George Patterson, Lieutenant Gouvernor cross section appearing at approximately 9.0 mm (NYSM of the State in 1848, not Paterson’s Gully. 13801); WW/WH = 100% by approximately 15 mm di- New material.–Most of the specimens reported here are ameter. WW/WH ratio continuing to decline, but at slower from three horizons in the Cashaqua Shale: (1) Parrish rate, and leveling off at approximately 35 mm diameter. At Limestone (approximately 60 specimens, mostly fragments); this stage and beyond, flanks sloping very gradually from (2) Randall Gully (Ontario County; Loc. 44/4; 10 speci- maximum WW on lower flanks to rounded venter (Text- mens); (3) Shurtleff Septarian Horizon (Livingston and On- figs 36E, 37A, C). Clarke’s (1898: pl. 4, fig. 16) illustration tario counties; approximately 50 specimens, all baritic of NYSM 3762 (traced as Text-fig. 36A), showing strong replacements). convergence of flanks toward venter, is misleading because Dimensions.–See Table 11. it is based on a specimen in which the body whorl is molded Diagnosis.–Moderately involute shell with cordate or in plaster (Pl. 9, Fig. 8). Flanks of mature specimens gently heart-shaped cross section. Internal whorls with prominent sloping to narrowly rounded venter (Text-figs 36B-C, G, lamellae (seen in baritic replacements) that become growth- 37A, C, Pl. 10, figs 1, 8. line lirae. Flanks of mature shell sloping gently from maxi- Suture distinctive, with large, rounded, projecting lateral mum whorl width (WW) on the lower flanks to narrowly saddle, acute lateral lobe, and broad umbilical saddle (Text- rounded venter. Suture with large projecting lateral saddle, figs 37E-H, Pls 9-10). acute lateral lobe, and broad umbilical saddle. Biconvex Elaborate ornamentation lamellae developing after ne- 126 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

Text-fig. 36. Cross sections of Manticoceras sinuosum sinuosum (Hall, 1843) from the Cashaqua Shale. A. NYSM 3762, cross section of a specimen from the Naples beds, Naples, Ontario County, traced from Clarke (1899a, b: pl. 4, fig. 16; body whorl of specimen modeled in plaster), 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 a specimen 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 section including portion of whorl beyond 36.2 mm diameter, from the Parrish Limestone in a tributary to Naples Creek (Loc. 46e/1) above NY Rte. 21, near Naples, Ontario County. F. X 3.6. G. X 1.4.

pionic constriction (Pl. 11, figs 5-6, 11-13); this feature re- based on numerous baritic replacements “from scattered cal- marked upon by Landman et al. (1996: 357-359). Clarke’s careous concretions” in the lower Naples beds between Ho- (1898, 1899a, b) detailed description of the shell develop- neoye and Conesus lakes; most probably came from the ment and ornamentation of Manticoceras pattersoni was Shurtleff Septarian Horizon of the upper Cashaqua Shale in HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 127

Text-fig. 37. Sutures and cross sections of Manticoceras sinuosum sinuosum (Hall, 1843) from the Cashaqua Shale. A-B. AMNH 5887/1, cross section and growth line based on the lectotype designated by House (1962), figured by Hall [1843: text-fig. 106(6)], and recorded as from Cashaqua 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 on a 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) as Goniatites patersoni Clarke, 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 a specimen from the Parrish Limestone, Rumpas Hill (Loc. 47a/1) near Naples, Ontario County, X 0.8. Scale = 5 mm (C). that area. Some of Clarke’s specimens are refigured here to approximately the end of the second whorl; at approxi- along with newly collected specimens from the Shurtleff mately this stage a shallow sinus (hyponomic sinus) develops Septarian Horizon. The distinctive ornamentation lamellae, on the venter and a slight sinus is seen on the mid-flank (Pl. which follow the smooth nepionic stage, gradually evolve 11, Figs 3-8, 11-13). The weakly sinuous, biconvex pattern into typical growth lines. Details of their development were persists to the largest diameters seen (Pl. 10, Figs 9-11, Pl. completely described by Clarke (1898, 1899a, 1889b: 53- 11, Figs 1-2). The slight umbilical salient and lateral sinus 54) but it is important to note that the major lamellae are on the mid-flank and the broad ventrolateral salient are dis- widely spaced and cross the shell with little or no curvature tinctive. Weak revolving lines on the lower flanks are seen at 128 B LEISO MRCNPLOTLG o 374 no. PALEONTOLOGY AMERICAN OF ULLETINS

Text-fig. 38. Graphs showing the ontogeny of Manticoceras sinuosum sinuosum (Hall, 1843). See Table 11 for biometric data. A. Based on measurements of the hypotypes from the Cashaqua Shale. Smaller specimen based on NYSM 3773, figured Clarke (1898: 41, fig. 1). Larger specimen based on NYSM 4762. B. Based on specimens from the Cashaqua Shale, NYSM 13800-13817. C. Based on specimens from the Shurtleff Gully Concretion Horizon, Cashaqua Shale, NYSM 13818-13835. HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 129

large diameters (Pl. 10, Fig. 10). The wrinkle layer (Pl. 15) tarian Horizon, between Conesus and Honeoye lakes (Liv- passes from the inside of the flanks of the body chamber ingston and Ontario counties; Locs 40-42). (ventral side) to the dorsal side where it rests on top of the Range.–Frasnian: Probeloceras and Prochorites Genozones preceding whorl; these ventral and dorsal wrinkle layers, re- UD I-E, F. Regional Zones of Probeloceras lutheri (19) and spectively, of House (1971) are clearly confluent. Prochorites alveolatus (20). Conodont Zones MN 5-6. Discussion.–Clarke (1898, 1899a, b) provided a detailed description of Manticoceras pattersoni (=M. sinuosum sinuo- Manticoceras sinuosum clausium n. ssp. sum) that was summarized with full synonymy by Miller Pl. 12, Figs 1-7, 11, Pl. 13, Figs 1-15; Text-figs 39A-I, (1938). Clarke treated M. sinuosum as the standard expres- 40A-B; Table 12 sion of the genus Manticoceras in New York. The other species of Manticoceras in New York were described in terms Etymology.–Latin clausium, a more enclosed place, refer- of “variations from this standard” often in terms of phases in ring to the tighter coiling in the early stages compared with ontogeny and geographical (ecological) distribution (Clarke, the nominate subspecies. 1898: 79-83, text-fig. 63); some are here assigned to other Type material.–Holotype NYSM 13839 (Pl. 12, Figs 3- species and to other genera such as Chutoceras, Koenenites, 4), Cashaqua Shale, Randall Gully (Ontario County; Loc. Sphaeromanticoceras, and Carinoceras. Comparatively little 44/3). attention was paid to the details of stratigraphic position of Material.–Seventy specimens from the lower part of the Hall’s and Clarke’s specimens and because of major miscor- Cashaqua Shale at Randall Gully (Loc. 44/3), 2.5 mi south relations (corrected following Chadwick, 1935b), the ranges of Bristol Center (Ontario County; 30 specimens), and of the species of Manticoceras in New York remained uncer- Whetstone Brook (Loc. 42/5), 2 mi west of Honeoye (On- tain (Clarke, 1898: text-fig. 64). tario County; 40 specimens, including baritic replacements). Detailed descriptions are lacking for many species of Diagnosis.–Similar to Manticoceras sinuosum sinuosum Manticoceras in general, which makes comparisons difficult. but shell more tightly coiled in early stages, lateral lobe less Among European species regarded (directly or indirectly) as acute and lower on flanks, and ornamentation lamellae less close to or synonymous with Manticoceras sinuosum are: M. prominent, more closely spaced. and becoming biconvex at intumescens (Beyrich, 1837), M. cordatum (G. & F. Sand- earlier stage. berger, 1850), M. lamed (G. & F. Sandberger, 1850), and Description.–Tighter coiling (lower UW/D) most appar- M. buchii (d’Archiac & de Verneuil, 1842). House (1962: ent between 3 and 10 mm diameter (Text-figs 39A-F); shell 259) considered M. sinuosum to be closer to M. cordatum development similar to that of Manticoceras sinuosum sinu- than to M. intumescens; both M. sinuosum and M. cordatum osum beyond 15 mm diameter. Lateral lobe (Pl. 12, Fig. 5, have somewhat flattened flanks that converge (but not Pl. 13, Fig. 12) still rounded and positioned just ventrad of steeply) toward the venter. M. intumescens has a more robust, umbilical shoulder at diameters at which it is acute and rounded, subquadrate whorl form (Wedekind, 1913: 52). higher on flank in M. sinuosum sinuosum; also less of umbil- North American comparable species are Manticoceras ical saddle seen than in M. sinuosum sinuosum. regulare Fenton & Fenton, 1924, M. cordiforme Miller, Juvenile ornamentation lamellae less ornate, becoming 1938, and M. cf. sinuosum House & Pedder, 1963. Mantic- smaller, more sinuous, and closely spaced earlier than in oceras cordiforme (= M. septentrionale Miller, 1938), from the nominate subspecies. Lateral and ventral sinuses developing Northwest Territories, Canada, and M. sinuosum have a sim- in second volution; growth lines biconvex by start of third ilar growth-line pattern, but M. cordiforme has a more robust volution (Pl. 12, Figs. 7, 11, Pl. 13, Figs. 1, 10). In M. sin- whorl form (House & Pedder, 1963: 521, text-fig. 7). The uosum sinuosum, weak biconvex growth lines develop later in shell development and whorl-form of M. cf. sinuosum from third volution (Pl. 11, Fig. 1). the Northwest Territories compares closely with M. sinuosum Discussion.–Manticoceras sinuosum clausium n. ssp. has but there are differences in details of the suture. only been found in the lower Cashaqua Shale at two locali- Distribution.–Cashaqua Shale (Sonyea Group), Lake ties in Ontario County. Specimens of Manticoceras from Erie (Erie County) to Keuka Lake (Steuben County). Man- scattered horizons in the Cashaqua generally show better ticoceras sinuosum sinuosum occurs in notable abundance in agreement with the common nominate subspecies. Varia- the Parrish Limestone in the vinicity of Naples in Ontario tions in tightness of coiling (UW/D) do not seem to follow and Yates counties (Locs 46, 46a-f, 47, 47a-f), in concretion a stratigraphic sequence. For example, specimens of M. s. horizons in Barnes Gully, Canandaigua Lake (Ontario sinuosum from the Shurtleff Septarian Horizon of the upper County; Loc. 44a/2), Randall Gully (Loc. 44/4), south of Cashaqua in Randall Gully (Loc. 44/4) are more similar to Bristol Center (Ontario County), and in the Shurtleff Sep- M. s. clausium n. ssp. in this feature than specimens from 130 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

Table 12. Biometric data for Manticoceras sinuosum clausium n. ssp. from the Cashaqua Shale. See Text-fig. 40 for graphs.

Specimen D WW WH UW WW/WH UW/D

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 0.18 NYSM 13848 27.5 11.0 14.0 5.0 0.79 0.18 17.0 8.0 8.7 3.7 0.92 0.22 ca. 10.3 5.2 4.7 3.0 1.11 0.29 6.6 3.6 2.9 2.1 1.24 0.32 4.3 2.4 1.7 1.5 1.41 0.35 3.0 1.5 1.0 1.3 1.50 0.43 2.1 1.0 0.67 — 1.49 — NYSM 13842 23.0 ca. 10.0 11.5 ca. 4.0 0.87 0.17 NYSM 13849 16.6 7.7 8.3 3.0 0.93 0.18 10.2 5.4 5.1 2.2 1.06 0.22 6.2 3.5 2.8 1.7 1.25 0.27 3.9 2.3 1.7 1.4 1.35 0.36 2.6 1.3 0.90 1.0 1.44 0.38 NYSM 13854 10.3 5.7 5.0 ca. 2.3 1.14 0.22 6.4 3.8 2.9 1.9 1.31 0.30 4.0 2.4 1.7 1.5 1.41 0.38 2.7 1.5 0.96 1.1 1.56 0.41 1.9 1.0 0.64 0.80 1.56 0.42

the intervening Parrish Limestone. Range.–Frasnian: Probeloceras Genozone UD I-E. Re- Manticoceras sinuosum clausium n. ssp. would seem a log- gional Zone of P. lutheri (19). Conodont Zone MN 5. ical ancestor of M. s. sinuosum given its less advanced suture (less acute and more rounded lateral lobe) and earlier strati- Manticoceras sinuosum tardum Clarke, 1898 graphic appearance in the lower Cashaqua Shale. It is also Pl. 9, Figs 2-3, Pl. 14, Figs 1-10; Text-figs 39J, 40C; tempting to place M. s. clausium n. ssp. in a phylogenetic Table 13 line between M. simulator from the underlying Genesee Group and M. s. sinuosum from the upper Cashaqua Shale. Manticoceras tardum Clarke, 1898, 1899a, b: 63-64, 81, pl. 1, fig. 13. Too little, however, is known about M. simulator; the holo- ?Manticoceras tardum Clarke, 1898, 1899a, b: pl. 6, fig. 31 (NYSM type (NYSM 3797) and only known specimen is from an 3804). unknown horizon and locality (probably in the upper Gene- Manticoceras Pattersoni Clarke, 1898, 1899a, b (pars): 45-62, pl. 2, see Group) near Ithaca (Tompkins County). Manticoceras fig. 3. simulator still has a rounded lateral lobe on the umbilical Manticoceras tardum. Miller, 1938 (pars): 121-123, pl. 24, fig. 6. shoulder at 36.5 mm diameter but the umbilical width at (?) Manticoceras tardum. Miller, 1938: pl. 24, fig. 7. this diameter is even larger than in M. s. sinuosum. Manticoceras sinuosum Miller, 1938 (pars): 106-115, pl. 19, fig. 1. Manticoceras sinuosum clausium n. ssp. could be more closely related to M. contractum from the Genundewa Lime- Type material.—NYSM 3804, a plastotype, the larger stone (middle Genesee Group); unfortunately two of the and surviving syntype figured by Clarke, is here designated three figured syntypes are small, and nothing is known of as lectotype. A specimen figured as Manticoceras Pattersoni the suture. The umbilicus of M. contractum is narrow and by Clarke (1898, 1899a, b: pl. 2, fig. 3; NYSM 3766) is in- the whorl form similar (Clarke, 1898, 1899a, b: text-fig. 44) cluded in the subspecies. but slightly more compressed; its growth lines are distinctly New material.–Ten specimens (mostly small baritic re- biconvex by the end of the second volution (Miller, 1938: placements) from the Cashaqua Shale, Briggs Gully (Loc. 82) as in M. s. clausium n. ssp., but the lamellae tend to clus- 43/3), Honeoye Lake. Bed 43/3 is believed to be the source ter in pairs, forming fascicles. of the smaller syntype figured by Clarke (1898, 1899a, b: pl. Distribution.–Lower Cashaqua Shale (Sonyea Group) at 1, fig. 13), which is lost (Miller, 1938: 123). Whetstone Brook, (Loc. 42/5) near Honeoye (Ontario Dimensions.–See Table 13. County), and Randall Gully (Loc. 44/3) near Bristol Center Discussion.–Clarke (1898, 1899a, b) described Mantico- (Ontario County). ceras tardum as a species with a “primitive” ornamentation in HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 131

Text-fig. 39. Sutures and cross sections of Manticoceras sinuosum clausium n. ssp. and M. s. tardum Clarke, 1898, from the lower Cashaqua Shale. A-I. M. s. clausium n. ssp., from Randall Gully (Loc. 44/3) near Bristol Center, Ontario County, unless otherwise stated. A. NYSM 13849, cross section, including portion of whorl beyond, at 16.6 mm diameter, X 1.5. B. NYSM 13851, cross section, X 1.5. C. NYSM 13852, 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 17.5 mm 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), Honeoye Lake, Ontario County, X 0.9. which simple distant ornamentation lamellae persist without ment of approximately three whorls; the larger of the syn- intercalation to the end of the third whorl and a much wider types (NYSM 3804) is a plastotype of four whorls. umbilicus and also broader venter than M. pattersoni (= M. The protoconch is similar to that of Manticoceras sinuo- sinuosum). The locality given for the syntypes was the lower sum, but appears to be more globular in shape, as the whorl Portage shales of Briggs Gully; Miller (1938: 123) stated that section is strongly reduced after the nepionic swelling (Pl.14, they came only from the Cashaqua Shale at that locality. figs 1, 3). The first few whorls are rather flattened dorsovent- The smaller of the syntypes (now lost) was a baritic replace- rally, producing a broad venter, and narrowly rounded flanks 132 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

Table 13. Biometric data for Manticoceras sinuosum tardum Clarke, 1898, from the Cashaqua Shale. See Text-fig. 40 for graph.

Specimen D WW WH UW WW/WH UW/D

NYSM 13870 ca. 82 28.0 41.0 14.3 0.68 0.17 ca. 53.5 —— 9.0 — 0.17 ca. 33 ca. 13.0 17.0 ca. 6.6 0.76 0.20 ca. 20.5 —— ca. 5.5 — 0.27 NYSM 13873 17.0 7.1 7.0 ca. 5.0 1.01 0.29 NYSM 13872 ca. 5.6 —— 2.4 — 0.43 4.1 2.0 1.3 1.9 1.54 0.46 ca. 2.9 —— 1.4 — 0.48 2.2 0.96 0.65 1.0 1.48 0.45 NYSM 13871 5.4 2.8 2.0 2.3 1.40 0.43 3.8 1.8 1.2 ca. 1.7 1.50 0.45 2.8 1.3 ca. 0.83 ca. 1.3 1.57 0.46 2.0 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

and slight dorsal impressment. specimen. This is apparently the case with Manticoceras ap- Simple distinct growth lamellae are still seen in NYSM primatum. Here the wide umbilicus and simple persistant 13879 (Pl. 14, Fig. 1) at the end of three whorls. Low trans- growth lamellae of the smallest syntype (NYSM 3731; also verse ribs, which cross the flanks and venter, are seen in in- a plastotype) compare closely with M. sinuosum tardum ternal molds (Pl. 14, Figs. 2, 4, 6-7); the ribs and intervening (NYSM 3804) figured on the same plate. NYSM 3731 is broad depressions are traversed by fine, closely spaced stria- from the Naples beds (Cashaqua?), at Griswold. Based on tions. The major growth lamellae of the shell wall (Pl. 14, the two larger “syntypes” from the Genundewa Limestone, Figs 4, 6-7) overlie the ribs and depressions. however, M. apprimatum is distinguished by its very narrow The umbilical widths of the smaller specimens are only umbilicus: “The umbilicus is very deep and narrow, closer slightly larger than in Manticoceras sinuosum sinuosum, and than in any other species, even more extreme than in Man- they compare with the smaller of Clarke’s syntypes. The ticoceras contractum.” (Clarke, 1898, 1899a, b: 62). whorl form and umbilical widths of the larger specimens, The new specimens are conspecific with the smaller of however, cannot be distinguished from the nominate sub- the figured specimens of Manticoceras sinuosum tardum, species at the same stage (Text-fig. 39J). based on their similar growth-line lamellae, degree of umbil- The suture is seen near the end of the second whorl, and ication, and rather broad, flattened venter in the early here the ventral lobe is trifid, as in Manticoceras sinuosum whorls. The presence of larger forms that are indistinguish- sinuosum; the suture and growth lines of NYSM 13843 at 17 able from M. s. sinuosum suggests that M. tardum is best re- mm diameter also cannot be distinguished from the nomi- garded as a subspecies of M. sinuosum. nate subspecies at the same stage. Distribution.–Lower Cashaqua Shale (Sonyea Group) at The most serious problem in determining Manticoceras Briggs Gully, Honeoye Lake (Ontario County; Loc. 43/3). sinuosum tardum concerns the degree of umbilication in the Range.–Frasnian: Probeloceras Genozone UD I-E. Re- two syntypes. Clarke (1898, 1899a, b: 63) noted: “The um- gional Zone of P. lutheri (19). Conodont Zone MN 5. bilication of M. tardum is much greater than that of M. Pat- tersoni and M. apprimatum. . . .The form and crosssection Manticoceras lamed (G. & F. Sandberger, 1850) of the whorl is distinct from both at parallel growth stages, Pl. 16, Figs 1-2, 9-10; Text-figs 41A-C, 42E; Table 14 being broader and flatter on the venter.” Miller (1938: 122) examined the larger syntype (NYSM 3804) and cited the Goniatites lamed var. complanatus G. & F.Sandberger 1850: 90, pl. following dimensions: D = ca. 11.5 mm; UW = 6.5 mm; 8, figs 5, 5a; Wedekind, 1913: pl. 6, fig. 3. Miller, 1932: 331; House & Ziegler 1977: 83, WH = 4 mm. The UW value is significantly greater than in Manticoceras lamed pl. 2, figs 15-17, 23-24. M. sinuosum sinosum and the specimens in hand. The UW Manticoceras (Manticoceras) cordatum (pars) Clausen, 1969: 130. of the smaller syntype figured by Clarke, however, compares with specimens of M. pattersoni of similar size on the same Type Material.–A lectotype was chosen and illustrated by plate. House [in House & Ziegler, 1977: pl. 2, figs 15-17, Berlin It is possible that NYSM 3804 came from a different Museum (HUM) c. 491] from among the specimens illus- horizon and is not the same species as the smaller figured OS ICGSE:LT EOINGONIATITES DEVONIAN LATE KIRCHGASSER: & HOUSE

Text-fig. 40. Graphs showing the ontogeny of the subspecies of Manticoceras sinuosum (Hall, 1843). See Tables 12 and 13 for biometric data. A-B. M. s. clausium n. ssp., based on measurements of specimens from the Cashaqua Shale. A. NYSM 13839 and 13848-13849. B. 13840-13847, 13854-13868. C. M. s. tardum, based on measurements of NYSM 13869-13874. 133 134 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

Table 14. Biometric data for Manticoceras lamed (G. & F. Sandberger, 1850) from the Rhinestreet and Angola shales.

Specimen D WW WH UW WW/WH UW/D

NYSM 12057 94.3 33.2 48.9 ca. 17.6 0.68 0.19 NYSM 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 0.22 NYSM 12065 10.9 5.1 5.6 2.5 0.91 0.23 NYSM 12064 10.0 5.0 ca. 4.8 — 1.04 — NYSM 12063 7.5 3.8 ca. 3.8 ca. 1.2 1.00 0.16 NYSM 12062 5.6 3.3 ca. 3.4 — 0.97 —

trated by the Sandberger brothers; another of their speci- mens was also figured (pl. 2, figs 23-24). The type material was 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 mudrock and crystalline calcite mostly as internal molds but in some with the shell as a crystalline calcite replacement. Phragmo- cones and incomplete body chambers are represented. Dimensions.–The dimensions of the lectotype [Berlin Museum (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 with relatively small umbilicus. Maximum whorl width close to rounded umbilical shoulder; flanks converging in smooth curve to well-rounded to narrowly rounded venter. Sutures as illustrated in Text-fig. 41, often showing no subdivision of mid-ventral saddle in early whorls, but later saddle be- coming divided and increasing markedly in prominence rel- Text-fig. 41. Sutures of Manticoceras, Sphaeromanticoceras, and Del- ative to lateral saddle, which shows ratio change from ca. phiceras from the West Falls Group in New York State. A-C. M. 54% at 19 mm diameter to ca. 98% at approximately 68 lamed (G. & F. Sandberger, 1850). A. NYSM 12057, suture at an mm diameter (Text-figs 41A-B). Growth lines not seen in estimated 68 mm diameter based on a specimen from Angola large specimens, but in middle whorls show broad, flat lat- Shale equivalents in Wolf Creek (Loc. 85), Castile, Wyoming County, X 0.7. B. NYSM 12058, suture at ca. 19 mm diameter eral sinus passing with slight convexity to ventrolateral based on a specimen from the upper Rhinestreet Shale in Johnson salient and back to well-rounded ventral sinus. Creek (Loc. 58/1), Wyoming County, X 1.9. C. NYSM 12059, su- Discussion.–The name Manticoceras lamed is given prece- ture at ca. 6.5 mm diameter based on a specimen from lower An- dence over M. cordatum and the latter considered as a sub- gola Shale equivalents at Varysburg (Loc. 79/4), Wyoming County, species of M. lamed (see House & Ziegler, 1977). A X 3.1. D-E. M. aff. lamed, NYSM 12060. D. Suture at ca. 11 mm lectotype for M. cordatum was designated by House (in diameter, based on a specimen from the Point Breeze Goniatite House & Ziegler, 1977: pl. 2, figs 13-14), which is now the Bed, lower Angola Shale, in Relyea Creek (Loc. 82/6), Wyoming lectotype of the subspecies, a stouter variety. One New York County, X 1.9. E. Growth line at ca. 17 mm diameter, based on specimen (NYSM 12057; Pl. 16, Figs 9-10) could be refer- same specimen as D. F. S. rhynchostomum (Clarke, 1898), NYSM 12061, suture based on an incomplete specimen from equivalents able to the stouter subspecies. of the upper Rhinestreet Shale, in the probable Relyea Creek Hori- These rather laterally compressed New York manticocer- zon, in Kennedy Gulf (Loc. 63/3), near Dale, Wyoming County, atids with an elegantly ovoid cross section are close to Man- X 0.7. G. D. cataphractum (Clarke, 1898), NYSM 3643, lectotype ticoceras sinuosum, and much material could be here designated, suture at 8 mm diameter, of a specimen recorded indistinguishable from that species. The lectotype of M. s. as from the Hanover Shale, Java, Wyoming County, X 6.4. HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 135

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 Shale in Johnson Creek (Loc. 58/1), Wyoming County, X 2. C. C. vagans (Clarke, 1898), NYSM 12072, from the lower Angola Shale in Hampton Brook (Loc. 74/3), Erie County, X 2. D. S. oxy (Clarke, 1897), NYSM 12036, from the Point Breeze Goniatite Bed, lower Angola Shale in Big Sister Creek (Loc. 73/6), Angola, Erie County, X 2. E. M. lamed (G. & F. Sandberger, 1850), NYSM 12058, from upper Rhinestreet Shale equivalents in Johnson Creek (Loc. 58/1), Wyoming County, X 1.9. F. M. sinuosum apprimatum (Clarke, 1898), NYSM 3732, the lectotype, from the Genundewa Limestone, Middlesex, Yates County, X 2. G. C. sororium (Clarke, 1898), NYSM 3803, new cross section thought to be a cotype figured by Clarke (1899a, b: 76, fig. 59), believed to be former NYSM 12310/6, probably from the lower Angola Shale, near Angola, Erie County, X 2. H-I. S. rhynchostomum. H. NYSM 12073, from the Point Breeze Goniatite Bed of the lower Angola Shale 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.

sinuosum, however, shows a growth line with scarcely a lat- More work is needed on the European material before all eral sinus at all; as described previously, early stages of Hall’s this can be resolved. species seem to divide the ventral median saddle earlier. Distribution.–Upper Rhinestreet Shale and Angola Shale This is a common Frasnian species in Europe, but unfor- (West Falls Group) and equivalents in Erie and Wyoming tunately none of the European successions have provided a counties. Specimens from the upper Rhinestreet Shale in- satisfactory time sequence of material. The Büdesheim spec- clude: Cazenovia Creek (Erie County; Loc. 54/1), 3196/23; imens described by Clausen (1969), are mostly very small, Varysburg (Wyoming County; Loc. 57/2), NYSM 12064, so that it is difficult to compare with the Manticoceras sinu- 12065, and 12067; Johnson Creek (Wyoming County; Loc. osum and M. aff. lamed material described here from New 58/1), NYSM 12058; Relyea Creek (Wyoming County; York. Manticoceras lamed is described from Adorf, Rhenish Loc. 60/4), 3193/5; Kennedy Gulf (Wyoming County; Loc. Schifergebirge (House & Ziegler, 1977: 83) only from the 63/3), 3234/2. nodulosum Zone (lower cordatum Zone), and there was a Specimens from the lower Angola Shale include: Hamp- preference for using a German species name rather than im- ton Brook (Erie County; Loc. 74/4), 3274/4; Cazenovia porting the American name M. sinuosum for that material. Creek (Erie County; Loc. 75/4), 3251/1a; Sheldon Creek 136 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

Table 15. Biometric data for Manticoceras aff. lamed (G. & F. Sandberger, 1850) from the Angola Shale.

Specimen D WW WH UW WW/WH UW/D

NYSM 12068 22.9 8.7 11.5 3.1 0.76 0.14 NYSM 12069 ca. 18.1 7.6 9.5 3.0 0.80 0.17 NYSM 12060 16.8 7.7 7.2 ca. 2.5 1.07 0.15 NYSM 12070 12.6 5.2 ca. 6.6 ca. 2.5 0.79 0.20 NYSM 12071 12.4 5.9 7.1 — 0.83 —

(Wyoming County; Loc. 78/4b), 3226/9; Varysburg MN 4. (Wyoming County; Loc. 79/4), NYSM 12059, 12063, 3214/4. Manticoceras aff. lamed (G. & F. Sandberger, 1850) Specimens from the upper Angola Shale include: Lake Pl. 16, Figs 3-8; Text-figs 41D-E; Table 15 Erie Shore (Farnham Creek) (Loc. 71/9a), NYSM 12066; Big Sister Creek (Erie County; Loc. 73/9), NYSM 12062; aff. Goniatites lamed var. complanatus G. & F. Sandberger 1850: Varysburg (Wyoming County; Loc. 79/10), 3215/12; Wolf 90, pl. 8, figs 5, 5a. Creek (Wyoming County; Loc. 85), NYSM 12057. Range.–Frasnian: ?Late Beloceras-Playfordites-Neoman- Material.–Seven specimens referred to below and two ticoceras Genozones UD-I, ?H-J. Regional Zone of ?Schinde- others all preserved in mudrock and crystalline calcite as wolfoceras chemungensis to Sphaeromanticoceras solid specimens often slightly crushed in the outer whorls rhynchostomum (?21c-22b). Conodont zones very uncertain, with shell preserved where present as recrystallized calcite, all ?MN 8-11. from the Point Breeze Goniatite Bed in the lower Angola Shale. Manticoceras lamed aff. cordatum Dimensions.–See Table 15. (G. & F. Sandberger, 1850) Description.–Subinvolute with maximum whorl width Text-figs 33K-L close to rounded umbilical shoulder with flanks converging flatly to ventrolateral area with slight concavity in outer part Material.–Three fragments (NYSM 16568, 3911/14, often emphasized by diagenetic crushing; rounded but 3911/18) of calcitic replacements from the Fossil Log Hori- abrupt turnover of flanks to flatly rounded, wide venter. Su- zon of the upper West River Shale (the Williamsburgh Bed ture (Text-fig. 41D) showing distinctive, low, divided ventral of Over et al., 1999, 2003), 3.4 m below the Middlesex saddle, broad lateral saddle, and wide, rounded lateral lobe Shale in Beards Creek, Leicester (Livingston County; Bed at 11 mm diameter. Growth lines weakly biconvex (Text- 18a/4b). fig. 41E; Pl. 16, Fig. 7) with broad, flat, slightly prosiradiate Discussion.–Assignment to the subspecies Manticoceras sinus on flanks, asymmetric ventrolateral salient, and broad, cordatum lineage (House & Ziegler, 1977) is based primarily rounded sinus on venter. Growth-line striae prominent (Pl. on whorl form. The shell is widest across the lower flanks, 16, Figs 3, 5, 7) and often bunched, simulating weak, fine and the flanks slope gently to a broad, rounded venter (in- ribbing on outer whorls (NYSM 12060 and 12068). cluding NYSM 16568 at ca. 9.3 mm diameter). In cross sec- Discussion.–As noted above, Manticoceras lamed is tion, they resemble the rather tubby M. contractum of the recorded from the upper Rhinestreet Shale to the upper An- Genundewa Limestone, but because the growth lines are not gola Shale, but not from the Point Breeze Goniatite Bed, preserved, comparison with the distinctive pattern in that which yields the richest fauna of the lower Angola Shale. species cannot be made. The form here described as M. aff. lamed is obviously part The suture of NYSM 16568, one-half whorl back from of the M. lamed lineage, but the differences are sufficiently 12.5 mm diameter, shows a large, broad lateral saddle and a clear cut to draw attention to them by the use of open broadly rounded lateral lobe. nomenclature because there is a greater stoutness to the Distribution.–Upper West River Shale (Genesee Group), whorl section and a more abrupt turnover from the flanks to Fossil Log Horizon (Williamsburgh Bed of Over et al., a wider rounded venter than in M. lamed, from which they 1999, 2003), Beards Creek, Leicester (Livingston County; also differ in lacking the elegant whorl outline of the longer Bed 18a/4b). ranging form. The whorl form approaches that of M. schell- Range.–Frasnian: Timanites Genozone UD I-C. Re- wieni Wedekind (1913: 65, pl. 5, figs 1-2) but the holotype gional Zone of Koenenites beckeri (17b). Conodont Zone of that species is noticeably more compressed (at D = 27 HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 137

mm, WW = 7.6 mm) and the flanks are rather flatter and and includes: C. acutum (G. & F. Sandberger, 1850: pl. 7, more parallel sided than the New York material. figs 1, 1a, 1b; 1851: 82); C. acutiforme (H. & G. Termier, Distribution.–Known only from the Point Breeze Goni- 1950: 53, 151, text-figs 13-14). Note that Manticoceras oxy atite Bed in the lower Angola Shale (West Falls Group) in is here referred to Sphaeromanticoceras because it is shown Wyoming County, at Relyea Creek (Loc. 82/6), NYSM that the discoidal form is a late stage development. 12060, 12068-12070, 3211/1a, and 3212/7, and Stony Oxyconic form is an iteratively repeated ammonoid Creek (Loc. 83/6), NYSM 12071. morphology (House, 1981a: 17) and there is no reason to Range.–Frasnian: Neomanticoceras Genozone UD I-J. suppose it arose only once from Manticoceras in the Ge- Regional Zone of Sphaeromanticoceras rhynchostomum (22b). phuroceratidae. Nevertheless, these two groups of species Conodont Zone ?MN 11. form a compact group. Possible derivations of it are forms such as M. inversum (Wedekind, 1913) and M. carinatum Manticoceras sp. (Beyrich, 1837), which are evolute in innermost whorls, Pl. 17, Fig. 4 oxyconic or carinate and sagittate in their middle whorls, but that revert to a typical M. lamed-cordatum type with Manticoceras accelerans Clarke, 1897: 53, nom. nud.; Clarke, 1898: rounded outline in outer whorls. 53; 1899a, b: 77, 82, text-figs 60, 63-64, pl. 6, fig. 10; Miller, Separation into two groups here is to emphasize the two 1938: 74, text-fig. 12, pl. 13, fig. 8. different patterns of growth-line form in mature individuals. In the second group, there is only a lateral salient, more lat- Type Material.–The only available syntype of Mantico- eral than ventrolateral in position. In the first group, there ceras accelerans, here designated lectotype, is NYSM 3730 are generally biconvex growth lines, but the original figures (formerly 12300/1). of Costamanticoceras koeneni (Holzapfel, 1882) show bicon- Description.–See accounts of Clarke (1898, 1899a, b: 77) vex growth lines in early whorls and then an increasing and Miller, (1938: 74). prominence of a broad ventrolateral salient (Holzapfel, Discussion.–We regard this crushed and unsatisfactory 1882: pl. 46, figs 4, 4b, 6) that suggests an intermediate po- lectotype as unusable for taxonomic purposes. However the sition (see Becker & House, 1993), and even the holotype generic assignment seems probable. of C. vagans described here shows a similar tendancy to a Distribution.–Reported to be from Cashaqua Shale broad and prominent ventrolateral salient although the (Sonyea Group) at Naples (Ontario County). growth lines as a whole are biconvex; both of these species Range.–Frasnian: Probeloceras or Prochorites Genozone are substantially smaller than specimens referred to the sec- UD I-E or F. Regional Zone of Probeloceras lutheri (19) or ond group. Prochorites alveolatus (20). Conodont Zone MN 5 or 6. Distribution.–North America, Europe, Russia, North Africa, and Australia. Genus CARINOCERAS Ljaschenko, 1957 Range.–Frasnian: Prochorites Genozone to Neomantico- Type species.–Carinoceras menneri Ljaschenko (1957: ceras Genozone UD I-F to J; possibly to Archoceras and 199), by original designation. Crickites Genozones (UD I-K to L). Diagnosis.–Gephuroceratids with oxyconic, compressed, and often keeled outer whorls; innermost whorls evolute; Carinoceras sororium (Clarke, 1898) middle whorls oxyconic to sagittate or rounded, subinvo- Pl. 18, Figs 3-4, 6-9; Text-fig. 42G lute. Reaching oxyconic form at early stage. Adult with deep, narrow umbilicus. Suture as in Manticoceras or, as in type Manticoceras sororium Clarke, 1898, 1899a, b: 75-76, 82, text-figs species, with incipient subdivision of ventral lobe in outer 56-59, pl. 4, figs 1-5; Frech, 1913: 24; Miller, 1938: 117-119, whorls. Shell surface smooth, growth lines biconvex or con- text-figs 23A-D, pl. 24, figs 1-5. vex. Manticoceras (Carinoceras) sororium. Clausen, 1971: 182, 198. Discussion.–Genus comprises two groups of species. The Carinoceras sororium. House & Kirchgasser, 1993: 276. first with biconvex growth lines includes: Carinoceras men- neri Ljashenko, 1957: 199 (pars), pl. 1, fig. 2; C. galeatum Type material.–Lectotype here selected as NYSM 3802 (Wedekind, 1913: 60, pl. 4, figs 3-4, text-fig. 8a); C. soror- (Pl. 18, Figs 8-9). Original specimens (NYSM 3798-3803) ium (Clarke, 1898, 1899a, b: 75, 82, pl. 4, figs 1-5, text- are gutta percha molds obtained after dissolution of calcitic figs 56-59, 63-64; cotypes NYSM 3800-3802); and C. replacements. This method of extraction shows the shell or- vagans (Clarke, 1898, 1899a, b: 78, 82, pl. 6, figs 11-12, nament superbly, but destroys other shell characteristics. text-figs 62-64; holotype NYSM 3805). Dimensions.–The dimensions of figured material of The second group appears to have convex growth lines Clarke (1899a, b) are as follows: NYSM 3798 (pl. 4, fig. 1), 138 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

Table 16. Biometric data for Carinoceras vagans (Clarke, 1898) Group from the Rhinestreet and Angola Shales. See Text-fig. 46A for graph. Specimen D WW WH UW WW/WH UW/D

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 0.19 NYSM 12044 31.5 ca. 12.3 ———— NYSM 12036 29.9 8.2 18.2 ca. 3.0 0.45 9.9 NYSM 12043 24.6 ca. 11.0 12.0 5.8 0.92 0.24 NYSM 12042 15.4 6.5 7.8 -- 0.83 — NYSM 12041 12.8 5.4 6.7 ca. 3.1 0.81 0.24 NYSM 12040 11.8 5.8 5.8 ca. 3.2 1.00 0.27 NYSM 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

D = 6 mm; NYSM 3799 (pl. 4, figs 2-3), D = 5.4 mm; tographically for the first time here (Pl. 23, Figs 13-14) and NYSM 3800 (pl. 4, fig. 4), D = 11.2 mm; NYSM 3801 (pl. a new preparation has been made of the suture (Text-fig. 4, fig. 5), D = 11.3 mm. 43A). Description.–Detailed descriptions were given by Clarke New Material.–Eleven specimens (including NYSM (1899a, b: 75-76, 82) and Miller (1938: 117-119) and need 12036-12042 and 12048) have been found in situ during not be repeated. The original material is partly reillustrated this work, all from the upper Rhinestreet Shale and Angola photographically here for the first time (Pl. 18, Figs 3-4, 6- Shale. The specimens are mostly small but preserved solid in 9). The technique of mold preparation leads to uncertainty mudstone often with growth lines and frequently with su- on the precise shape of outer whorls. tures shown. Discussion.—This species has not been identified in our Dimensions.–See Table 16 and Text-fig. 46A. collections. Clarke, however, illustrated distinctive early Description.–Innermost couple of volutions evolute; stages. It is considered that it might represent early stages whorl section rounded, depressed; by 6 mm diameter similar to Carinoceras vagans but an adult with rounded (NYSM 12040) carinate form of adult already seen (Text- rather than acute venters. fig. 43E) and WH = WW at approximately 12 mm diameter Distribution.–NYSM 3798-3803 are recorded from “the (NYSM 12040); thereafter whorl height increasing relatively calcareous concretionary masses in the shales on Big Sister rapidly (Text-fig. 46A). Maximum whorl width at 18 mm Creek, Erie County” (Clarke, 1899c: 106-107) and all spec- diameter close to umbilical shoulder and flanks converge imens known to Clarke were from the vicinity of Angola flatly, but with slight convexity, to acute venter. At approx- (Loc. 73). The source of the material is probably all from imately 20 mm diameter, whorls have maximum width close within the lower Angola Shale. Clarke mentioned Farnham to umbilicus and flanks slope convexly and evenly to suba- Creek as another locality (Locs 71-72). cute venter. By 27 mm diameter (NYSM 12036), trend con- Distribution.–Lower Angola Shale (West Falls Group; tinuing with venter so sharply acute to feel knife-edged; Erie County). suture near this diameter illustrated in Text-fig. 44D. Larger Range.–Frasnian: Neomanticoceras Genozone UD I-J. specimens (NYSM 12044) much stouter in cross section but still galeate. Sutures of earliest whorls unknown. Growth Carinoceras vagans (Clarke, 1898) lines biconvex at all stages seen, with shallow latero-umbil- Pl. 23, Figs 5-16; Text-figs 43A, D-E, 44D, 46A; Table 16 ical salient essentially on shoulder and increasingly broad and prominent ventrolateral salient (Text-fig. 43D); growth Manticoceras vagans Clarke, 1898, 1899a, b: 78-79, 82, text-fig. lines not seen above 23 mm diameter. 62, pl. 6, figs 11-12; Frech, 1913: 24; Miller, 1938: pl. 15, figs Discussion.–The stratigraphically youngest specimens 11-14; House, 1962: 258-259. known, from above the level of the Point Breeze Goniatite Manticoceras (Carinoceras) vagans. Clausen 1971: 182, 198. Carinoceras vagans. House & Kirchgasser, 1993: 276. Bed (NYSM 12043-12045), are determined as Carinoceras Carinoceras oxy House & Kirchgasser, 1993: figs 4E-F. aff. vagans on account of the rather more robust whorl sec- tion and the more forward position of the lateral lobe. Type Material.–Only one specimen, the holotype The group of “species” Carinoceras menneri, C. galeatum, (NYSM 3805), was known to Clarke (1898,1899a, b) and C. vagans, C. acutum, and C. acutiforme share much in com- to Miller (1938), who each described it. It is illustrated pho- mon and there is a case to be made for preserving the names HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 139

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 recorded as 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, suture at ca. 27 mm diameter based on a specimen from Angola Shale equivalents at Varysburg (Loc. 79/10), Wyoming County, X 4.5. C. NYSM 12044, suture at ca. 21 mm diameter based on a specimen from above the Point Breeze Goniatite Bed in Angola Shale equivalents at Glade Creek (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), Wyoming County, X 4.5. E. NYSM 12040, cross section of a juvenile at 13.9 mm diameter reconstructed from a specimen from the upper Rhinestreet Shale equivalents on Johnson Creek (Loc. 58/1), Wyoming County, X 4.1. F-G. S. rhynchostomum, NYSM 12035, suture at WH = 64 mm with 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. 140 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

est to C. galeatum and there is little doubt they should be re- garded as synonyms, C. vagans having priority. But the latter is here shown to modify into forms with very sharp oxyconic venters. Carinoceras acutum would seem to be close to C. acutiforme but to differ only a little from C. vagans insofar as 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 for carination or adult growth line convexity in the former. It is probable, as others have remarked, that Carinoceras galeatum is a junior synonym of this group, and the larger of the measurements given by Wedekind (1913: 60), that are plotted 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 at present unknown, and speculation on relationships with Eu- ropean forms is best deferred until a detailed study has been undertaken. Text-fig. 44. Sutures and growth line of Sphaeromanticoceras spp. Distribution.–Upper Rhinestreet Shale to middle Angola from the West Falls Group in New York and Carinoceras spp. from Shale (West Falls Group) in Erie and Wyoming counties and New York and Belgium. A. S. oxy (Clarke, 1897), NYSM 3747, probably the equivalent interval (Grimes Siltstone to Nunda growth line based on one of Clarke's specimens recorded as from Sandstone (High Point Sandstone) in the Naples area (On- the "Portage sandstone on East Hill, Naples," X 0.4. B. C. acutum tario County). The stratigraphic horizon of the holotype (G. & F. Sandberger, 1850), IG 8633, suture at 78 mm whorl height and reversed for comparison, of a specimen determined by (NYSM 3805) is unknown but the specimen almost cer- Matern from the Schistes de Matagne (F3) in the railway cutting tainly came from the interval from the Grimes Siltstone to between Nismes and Mariembourg, Belgium, X 0.4. C. S. oxy, the Nunda Sandstone of the West Falls Group (Text-fig. 3). NYSM 3746, the last suture, at 137 mm diameter and reversed Clarke (1899a, b: 79) stated that the specimen came from for comparison, of another of Clarke's specimens, from 45.7 m “. . . a loose block of sandstone among the Portage outcrops (150 ft) "below the Portage Sandstone on the Genesee River" (Loc. in the town of Naples, . . . associated with [the brachiopods] 86), Wyoming and Livingston counties, X 0.4. D. C. vagans Productella speciosa [Hall, 1867], Leptostrophia mucronata (Clarke, 1898), NYSM 12036, suture at 23 mm diameter based on [Hall, 1867] and Amboecoelia umbonata [Conrad, 1839].” a specimen from the Angola Shale (Point Breeze Goniatite Bed) on Miller (1938: 125) stated that Chadwick “. . . has recently Big Sister Creek (Loc. 73/6), Angola, Erie County, X 0.8. E. S. oxy, NYSM 3748, lectotype here designated, portion of the ventral given its horizon as the Grimes sandstone member of the suture on one of Clarke's specimens labeled as from Naples, X 0.4. Chemung formation” but gave no reference. Chadwick F. S. aff. oxy, NYSM 12034, portion of a mature suture at 135 (1935a: 320-322) listed Carinoceras vagans, C. sororium, and mm whorl height, reversed for comparison, showing obliquely flat- Spheromanticoceras oxy as confined to the Chemung group tened crest to the ventrolateral saddle, based on a specimen from (the base of which at Naples is the Grimes) but of the three the Relyea Creek Horizon of the Rhinestreet Shale on Relyea associated brachiopods, Chadwick (1935a) listed only P.spe- Creek (Loc. 60/3), Wyoming County, X 0.4. ciosa as a “transient” brachiopod extending into the post- Chemung sequence. Clarke & Luther (1904) did not list merely as subspecies within the oldest available name, C. any of the three brachiopods in their faunal lists of the acutum. Comparisons are aggravated by the lack of much Grimes or succeeding West Hill Flags and Sandstone (in statistical data, let alone stratigraphical data, on the forms which S. oxy is listed). Their faunal list for the High Point concerned. Those data available have been plotted in Text- Sandstone at the top of the Naples succession includes P. fig. 46C. These are based wholly on the suite of type speci- speciosa and A. umbonata. mens in each case except that Büdesheim data from Clausen We follow Pepper et al. (1956) in equating the High (1969) are also added for early whorls. Clausen assigned the Point Sandstone at Naples with the Nunda Sandstone (Text- numbered specimens as follows: C. galeatum, 36-40; C. aff. fig. 3) which puts the Nunda as an upper limit to the pos- vagans, 71-75; C. acutiforme, 76-80; assignment of these nu- sible range of the type of Carinoceras vagans. clei, however, is open to considerable doubt. The stratigraphically lowest specimens in our collections Considering shell form alone, Carinoceras vagans is clos- are upper Rhinestreet Shale: two specimens (NYSM 12040 HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 141

and 12042) from the fourth microcyclothem below the top I-K, L. of the Rhinestreet on Johnson Creek (Wyoming County; Loc. 58/1), and four specimens from the overlying third mi- Delphiceras cataphractum (Clarke, 1898) crocyclothem down (NYSM 12048), NYSM 3200/29 from Pl. 19, Figs 1-9; Text-fig. 41G the Relyea Creek Horizon (Loc. 60/3), NYSM 3193 from Relyea Creek (Wyoming County; Loc. 60/4), and NYSM Gephyroceras cataphractum Clarke, 1898, 1899a, b: 87-89, text- 3297/19 from Varysburg (Wyoming County; Loc. 57/2). figs 66-67, pl. 6, figs 3-9. . Frech, 1913: 25; Miller, 1938: 77-80; The specimens from the Angola Shale include NYSM Manticoceras cataphractum House, 1962: 258; House & Kirchgasser, 1993: 275, 279. 3248/2, from the fourth microcyclothem up from the base, Delphinites cataphractum. Becker & House, 1993: 114, pl. 2, figs on Cazenovia Creek (Erie County; Loc. 75/4), and NYSM 4-6. 12037 from the sixth microcyclothem, just below the Point Delphiceras cataphractum. Becker & House, 2000: 130, 140. Breeze Goniatite Bed on Hampton Brook (Erie County; Loc. 74/6a). The best fauna (NYSM 12038, 12039, 12041, Type material.–Seven original cotypes of Clarke (NYSM and 3229/6, 31) is from the Point Breeze Goniatite Bed it- 3638-3644; formerly 12160/1-7). Lectotype here chosen as self on Cazenovia Creek (Loc. 75/6). NYSM 12036 is from NYSM 3643 (figured by Miller, 1938: pl. 13, fig. 1 and the Point Breeze Goniatite Bed on Big Sister Creek, Angola herein Pl. 19, Fig. 1 and Text-fig. 41G). These fall into two (Erie County; Loc. 73/6). The stratigraphically highest spec- groups: NYSM 3638 and 3643 are limestone internal imens, referred to Carinoceras aff. vegans (Pl. 23, Figs 1-4; molds; NYSM 3639-3642 and 3644 are baritic shell replace- Text-figs 43B-C) are from middle levels of the Angola Shale, ments. NYSM 12043 and 12045 from 10 m above the Point Breeze Description.–Detailed descriptions have been given by Goniatite Bed at Varysburg (Wyoming County; Loc. 79- Clarke (1899a, b: 87-89) and Miller (1938: 77-80) and need 80/10), and a higher specimen (NYSM 12044) from Glade not be repeated here. The prominent constrictions are diag- Creek (Wyoming County; Loc. 77/1). nostic. Range.–Frasnian. Playfordites to Neomantioceras Geno- Distribution.–Lower Hanover Shale (West Falls Group; zones UD I-I-J. Regional zones of P. cf. tripartitus (22a) and Wyoming and Chautauqua counties). The cotypes are Spheromanticoceras rhynchostomum (22b). Conodont Zone recorded as from calcareous concretions near the village of MN 11-?12. Java (Wyoming County), at levels subsequently referred to the Hanover Shale; the horizon at Java is in Beaver Meadow Genus DELPHICERAS Becker & House, 2000 Creek (Bed 92/2). We have only found specimens in the Type species.–Manticoceras unduloconstrictum Miller, nodules in the lowermost microcycles in the Hanover Shale, 1938, by original designation (Becker & House, 1993: 114); including localities close to Lake Erie Shore (Text-fig. 16), holotype illustrated by Becker & House (1993: pl. 2, fig. 7) such as Walnut Creek (Loc. 89a/2-4), south of Silver Creek and herein (Pl. 18, Fig. 10). (Chautauqua County). Diagnosis:–Small-sized gephuroceratids, moderately evo- Range.–Frasnian: Archoceras Genozone UD I-?K. Re- lute to subinvolute throughout ontogeny with whorl height gional Zone of Delphiceras cataphractum (23). MN 12. rather rapidly expanding. Rounded whorl section with ven- ter rounded or slightly flattened. Periodic internal shell Genus PLAYFORDITES Becker, House & thickenings forming constrictions on the internal molds. Kirchgasser, 1993 Growth lines rectiradiate to biconvex. Type species.–Goniatites lamed var. tripartitus G. & F. Included species.–Manticoceras unduloconstrictum Miller, Sandberger, 1850. Original material lost; neotype chosen by 1938, and Gephyroceras cataphractum Clarke, 1898. Becker & House (1993: 121) is the specimen figured by Discussion.–The generic name originally proposed for House (in House & Ziegler, 1977: pl. 3, figs 13-15), and this group was Delphinites Becker & House, 1993: 114, but by Becker et al. (1993: fig. 4, O, P; Univ. Marburg Coll., that proved to be preoccupied by Delphinites Sayn, 1901. Mbg. 2594a). Delphiceras was proposed as a replacement name (Becker & Diagnosis.–Slender gephuroceratids with discoidal shell House, 2000: 140) and hence takes the same type species in middle whorls with spiral thickenings on either side of automatically. venter or on flanks, which are not shown on external surface Distribution.–North America (Indiana and New York) of shell. and Europe (England, Belgium, Germany) (Becker & Included species.–Playfordites tripartitus (G. & F. Sand- House, 1993). berger, 1850). Range.–Frasnian. Archoceras to Crickites Genozones UD Distribution.–North America (New York), Europe (Ger- 142 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

Table 17. Biometric data for Playfordites cf. tripartitus (G. & F. Sandberger, 1850) from the Rhinestreet Shale.

Specimen D WW WH UW WW/WH UW/D

NYSM 12074 9.2 4.4 4.4 ca. 2.0 1.00 0.22 5.6 3.1 2.6 1.5 1.19 0.27 3.7 2.1 1.5 1.2 1.40 0.32 NYSM 12075 8.2 4.3 ————

many), and Western Australia. Diagnosis.–Small to very large gephuroceratids with Range.–Frasnian: Playfordites to Neomanticoceras Geno- widely umbilicate earliest whorls followed by stages with a zones UD I-I-J. very depressed whorl section, often banana-shaped. Outer whorls of large forms laterally compressed and discoidal. Su- Playfordites cf. tripartitus (G. & F. Sandberger, 1850) ture with mid-lateral saddle very high, symmetric to asym- Text-fig. 42A; Table 17 metric with steep to overhanging dorsad face. Growth lines biconvex in early stages and small forms to subbiconvex to Playfordites cf. tripartitus. House & Kirchgasser 1993: 276. subconvex in very large discoidal specimens. Wrinkle layer characteristically with much finer striae (in type species) Material.–Two small specimens only (NYSM 12074- than in Manticoceras. 12075), both preserved in mudrock and calcite replacement Included species.–Goniatites affinis Steininger (1849, nom. as internal molds showing part of the body chamber and nud.; 1853: 42, pl. 1, figs 4, 4c); Manticoceras hunanense Xu, phragmocone. 1977; M. kwangsiense (Chao, 1956: 106, pl. 1, figs 19-22; Dimensions.–See Table 17. = affine fide Becker & House, 1993); M. kweipingense Chao, Description.–Subinvolute, with open umbilicus and well- 1956 (? = orbiculus fide Becker in litt.); Goniatites lamed var. rounded shoulders (Text-fig. 42A) with maximum width latidorsalis G. & F.Sandberger (1850: 90, pl. 8, fig. 8; = affi- close to shoulder and flanks converging convexly toward nis fide Clausen, 1969); Ammonites orbiculus Beyrich (1837: rounded venter. In NYSM 12074, ventrolateral furrows are 36, pl. 2, figs 4a-b); M. oxy Clarke, 1897; A. primordialis well developed with arched band between. In NYSM von Buch (1832: 168, pl. 1, figs 15-17; possible senior syn- 12075, in addition to ventrolateral groove, is another but onym of affinis); M. rhynchostomum Clarke, 1898; Sphaero- slighter spiral concavity on outer flanks, umbilicad of fur- manticoceras rickardi House & Kirchgasser, 1993. row. Growth lines forming slight salient on umbilical shoul- Discussion.–Although it is accepted that Clausen’s taxo- der and sinus, which centers low on flanks; lines then nomic separation of this group is useful, so many of the type sweeping well forward to ventrolateral salient on umbilical specimens of species that he included within it are poorly side of furrow; ventral sinus deep, rounded. Suture (NYSM known and badly described that there is more than usual 12074) showing small, undivided ventral saddle, broadly uncertainty on the assignments given here. A further prob- rounded and nearly symmetrical lateral saddle, and broad lem is that Sphaeromanticoceras affine (Steininger, 1849: 2- and rounded lateral lobe (L) low on flanks. Ventral wrinkle 3, 42, figs 4, 4a), the type species, is very small layer showing as fine, discontinuous striae on internal mold, (approximately 11 mm in diameter), whereas late Frasnian which are directed in convex rursiradiate course across species can reach over a third of a meter in diameter. flanks. Distribution.–Widely distributed in North America, Eu- Discussion.–This is a very rare form and so, unfortu- rope, Asia, and Australia. nately, comparison cannot be made with the varied develop- Range.–Frasnian: Prochorites to Crickites Genozones UD ments of specimens from the Canning Basin, Australia. I-F-L. Distribution.–Upper Rhinestreet Shale to Lower Angola Shale (West Falls Group) in Wyoming County. NYSM Sphaeromanticoceras rhynchostomum (Clarke, 1898) 12074 from the Upper Rhinestreet Shale, Johnson Creek Pl. 20, Figs 1-12, Pl. 21, Figs 1-12, Pl. 22, Figs 1-9; Text-figs (Loc. 58/1) and NYSM 12075 from the Point Breeze Goni- 41F, 42B, H, I, 43F-G, 47; Table 18 atite Bed of the Angola Shale, Relyea Creek (Loc. 82/6). Manticoceras rhynchostoma Clarke, 1898, 1899a, b: 65-69, 81, pl. Genus SPHAEROMANTICOCERAS Clausen, 1971 4, figs 6-13, pl. 5, fig. 1, text-figs 30-43; 1904: 356, 361, 365; Type species.–Goniatites affinis Steininger, 1849, by orig- Frech, 1913: 24; Miller, 1938: 99-102, pl. 14, fig. 19, pl. 25, inal designation (Clausen, 1971: 197). figs 1-8, pl. 26, figs 1-2, text-figs 18A-L; Clausen, 1969: 146. HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 143

Table 18. Biometric data for Sphaeromanticoceras rhynchostomum (Clarke, 1898) from the upper Rhinestreet and lower Angola shales. See Text-fig. 47 for graphs.

Specimen D WW WH UW WW/WH UW/D

NYSM 3796 94 32 52 ca. 13 0.62 0.13 (lectotype) NYSM 12132 27.9 16.0 15.7 7.0 1.02 0.25 NYSM 12080 31.1 15.3 15.0 8.6 1.02 0.28 NYSM 12085 17.2 10.5 7.4 5.4 1.42 0.31 11.3 7.1 4.5 4.0 1.58 0.35 7.9 5.0 2.9 3.1 1.72 0.39 5.4 3.3 1.8 2.2 1.83 0.41 4.0 2.3 1.2 1.8 1.92 0.45 2.9 1.6 0.9 1.3 1.78 0.45 2.2 1.2 0.7 0.9 1.71 0.41 1.6 0.8 0.5 0.7 1.60 0.44 NYSM 12088a 13.2 8.3 5.7 4.2 1.46 0.45 8.6 5.9 3.2 3.4 1.84 0.40 6.1 4.2 1.9 2.8 2.21 0.34 4.6 2.8 1.3 2.0 2.15 0.43 3.3 1.9 1.1 1.5 1.73 0.45 2.5 1.3 0.7 1.1 1.86 0.44

Sphaeromanticoceras rhynchostomum. House & Kirchgasser, 1993: (Loc. 74/6a); NYSM 12078 and 12079, Hampton Brook 275-276, figs 4I-J. (Loc. 74/6b) in Erie County; and NYSM 12091/1-4, Cazenovia Creek (Loc. 75/6); NYSM 12092, Johnson Creek Type material.–Cotypes are NYSM 3784-3796, and of (Loc. 81/6); NYSM 12093/1-23, Relyea Creek (Loc. 82/6); these, NYSM 3796 is here designated as lectotype (figured and coll. #3205, Stony Creek (Loc. 83/6a), in Wyoming by Clarke, 1898, 1899a, b: 68, text-fig. 43, and herein, Pl. County. 22, Figs 6-7, Text-fig. 47A). All are reported from Big Sister From higher in the equivalents of the Angola Shale, Creek (Erie County), and probably from the Angola Shale Sphaeromanticoceras rhynchostomum (or S. oxy) can be repre- at Angola (Erie County; Loc. 73). sented in the vicinity of the Nunda Sandstone at Varysburg New Material.–One-hundred fifteen specimens (includ- (Gassman Road) (Loc. 80/14) and at Wolf Creek (Loc. 85) ing NYSM 12035, 12061, and 12080-12093. Of these, 40 below Castile in Wyoming County. are from Rhinestreet Shale, 15 from the Angola Shale below Dimensions.–Dimensions are given in Table 18. Accom- the Point Breeze Goniatite Bed, 43 from the Point Breeze panying graphs separate type specimens (Text-fig. 47A), and Goniatite Bed or adjacent beds (NYSM 12087-12093), and specimens from the Rhinestreet (Text-fig. 47B) and Angola 7 from the Angola Shale equivalents above the Point Breeze shales (Text-fig. 47C). Goniatite Bed. Description.–Full descriptions were given by Clarke From the Rhinestreet Shale in the Relyea Creek Horizon (1898, 1899a, b) and repeated by Miller (1938). Some ma- or nearly correlative horizons: NYSM 12084/1-4, Cazenovia terial is in the form of baritic replacements and some type Creek (Erie County; Loc. 54/1); NYSM 12082, Varysburg material consists of gutta percha casts. The bulk of the new (Loc. 57/1); NYSM 12085, Johnson Creek (Loc. 58/1); sev- material is preserved as calcitic replacements with calcitic or eral specimens in collection 3200, Relyea Creek (Loc. 60/3); mudrock infills of phragmocone chambers, whereas the NYSM 12080/1-6, Stony Creek (Loc. 61/1); and NYSM body chambers are mostly preserved in calcareous mudrock. 12081/1, 2, Kennedy Gulf (Wyoming County; Loc. 63/3). Protoconch approximately 0.8 mm in diameter and Included is a fine collection (NYSM 6680/A-H, J-Q, and maximum width. (That the protoconch is wider than the 12132) by L. V. Rickard from the vicinity of the Relyea first whorl is clear from NYSM 12079; Pl. 20, Fig. 4.) No Creek Horizon at Relyea Creek (Loc. 60). lirae on the protoconch (ammonitella), which has a diameter From the Angola Shale, the best material is from the of approximately 1.1 mm. (An eccentrically placed am- Point Breeze Goniatite Bed: NYSM 12087/1-4, Point monitella is shown in Pl. 22, Fig. 9.) Breeze (Lake Erie Shore) (Loc. 72/6); NYSM 12088/1-3, Whorl following ammonitella widely umbilicate, with Big Sister Creek, Angola (Loc. 73/6); NYSM 12035, 12089, periodic raised lirae, numbering approximately 17 per whorl and 12090/1-6, bed below the Point Breeze Goniatite Bed and almost rectiradiate across flanks and venter (Pl. 20, Figs 144 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

Table 19. Biometric data for Sphaeromanticoceras oxy (Clarke, 1897) from the Rhinestreet and Angola shales. See Text-fig. 46 B for graph.

Specimen D WW WH UW WW/WH UW/D

NYSM 3746 201 — 105 24 — 0.12 (cotype) NYSM 3747 — ca. 47 ca. 150 —— — (cotype) NYSM 12034 355 — 177 ca. 32 — 0.09 NYSM 16536 299 ca. 100 162 39 0.61 0.13 176 ca. 70 75 29 0.93 0.16 147.7 52.4 81.2 19.3 0.65 0.13 101 47 50 23 0.94 0.23 87.4 31.8 46.0 15.2 0.69 0.17 61 33 28 16 1.18 0.26 48.8 23.1 24.1 11.8 0.96 0.24 29.7 19.2 13.6 7.4 1.41 0.25 19.3 11.4 7.7 6.2 1.48 0.32 12.9 7.8 5.3 4.6 1.47 0.36 8.7 5.2 3.0 3.5 1.73 0.40 6.8 4.3 2.2 2.5 1.95 0.37 5.0 3.4 1.5 1.9 2.27 0.38 NYSM 12046 ca. 137 — ca. 74 ca. 13 — 0.06 NYSM E-355 125 20 74 ca. 8.0 0.27 0.10 NYSM 12047 16.6 6.2 8.1 2.8 0.77 0.17

2-3). Whorl width increasing progressively to produce tubby these within S. rhynchostomum as is discussed below. shells by 10 mm diameter (Pl. 21, Figs 1-12, Pl. 22, Fig. 9; Distribution.–Upper Rhinestreet Shale and Angola Shale Text-figs 42B, H-I). Thereafter, WW/WH ratio declining (West Falls Group) in Erie and Wyoming counties. Com- (Text-figs 47A-C), reversing earlier pattern; change can mon in Relyea Creek Horizon of the upper Rhinestreet occur earlier in Rhinestreet than in Angola material. During Shale in Wyoming and Erie counties and the Point Breeze this stage, pattern of earlier lirae continuing but with gradual Goniatite Bed of the lower Angola Shale in Erie County. onset of slight biconvexity, increasing in frequency to ap- Range.–Frasnian: Playfordites and Neomanticoceras Geno- proximately 40 per whorl, with stronger development on zones UD I-I to J. Regional Zone of P. cf. tripartitus (22a) umbilical wall and lower flanks. Thereafter ornament pro- to Sphaeromanticoceras rhynchostomum (22b). Conodont gressively lost. Early lirae very dissimilar to virtual flares in Zone MN 11-?12. early stages of Manticoceras sinuosum Group. Subsequent growth as described by Clarke and Miller. Sphaeromanticoceras oxy (Clarke, 1897) Lectotype (Pl. 22, Figs 6-7; Text-fig. 47A) shows well the Text-figs 44A, C, E, 45, 46B; Table 19 gradual change in WW/WH ratio and how shells above ap- proximately 35 mm diameter become laterally compressed. Manticoceras oxy Clarke, 1897: 53; 1898, 1899a, b: 53, 77-79, Because of preservation, sutural ontogeny of earliest 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. stages not determined. Lectotype with relatively high me- 17, fig. 1, pl. 18, figs 1-3. dian saddle (MSh/ELh = 0.75 at 85 mm diameter) at which Carinoceras oxy. House & Kirchgasser, 1993: 276. point, for slightly asymmetric lateral saddle, LSw/LSh = 1.0. Lobe low on flanks (L) already as sharp as at 20 mm in some Type material.–The cotypes have been described and il- specimens. Note that in NYSM 12035, at WH = 61 mm lustrated by Clarke and by Miller and comprise the follow- (Text-figs 43F-G), the median saddle is very high (MSh/LSh ing figured by Clarke, 1898, 1899a,b: pl. 2, figs 5-6, = 1.14) and the low lateral lobe (L) very sharp. (NYSM 3746, a fine but crushed specimen), pl. 3, fig. 1 Discussion.–It is the characteristic tubby form in stages to (NYSM 3747, growth line illustrated in Text-fig. 44A), pl. approximately 20 mm diameter and the distinctive early li- 3, figs 2-3 (NYSM 3748, lectotype here designated, suture rate ornament that distinguishes this species. It is clear that refigured in Text-fig. 44E) and text-fig. 61 (NYSM 12305/4, large specimens become increasingly laterally compressed another large crushed specimen in which the whorl outline and discus shaped. End forms of this line are here placed in cannot be determined). None of these large and impressive Sphaeromanticoceras oxy. A case could be made for including specimens shows inner whorls, and the largest specimens HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 145

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, Relyea Creek (Loc 60/3), Wyoming County. C-D. NYSM 16570, side view and partial cross section of a specimen from a loose block, collected by B. Oldfield, of a shell-rich channel fill, upper West Falls Group (Nunda or Wiscoy Sandstone equivalents), at Loc. 87a, US Rte. I-390 in Steuben County, southeast of Dansville, Livingston County. Scales = 5 cm. 146 B LEISO MRCNPLOTLG o 374 no. PALEONTOLOGY AMERICAN OF ULLETINS

Text-fig. 46. Graphs showing the ontogeny of Carinoceras and Sphaeromanticoceras. A. C. vagans (Clarke, 1898) Group, based on material from the Rhinestreet and Angola shales; see Table 16 for biometric data. B. S. oxy (Clarke, 1897) [as C. oxy], based on material from the Rhinestreet and Angola shales; see Table 19 for biometric data. C. C. spp., based on material from Germany and Russia as indicated. OS ICGSE:LT EOINGONIATITES DEVONIAN LATE KIRCHGASSER: & HOUSE

Text-fig. 47. Graphs showing the ontogeny of Sphaeromanticoceras rhynchostomum (Clarke, 1898) from the Rhinestreet and Angola shales of western New York State; see Table 18 for selected biometric data. A. Based on cross sections (corrected for size) of NYSM 3796, the lectotype here designated, figured by Clarke (1899a, b: text-fig. 43) and NYSM 3794 (figured by Clarke, 1899a, b: text-fig. 41). Both probably from the lower Angola Shale from either Big Sister Creek or Angola, Erie County. B. Data from 23 specimens from the upper Rhinestreet Shale (Locs. 54/1, 57/1, 57/2, 58/1, 60/3, 61/1, and 63/3), all within 3-10 m (10-33 ft) below the top at localities between Cazenovia Creek, Erie County, and Kennedy Gulf, Wyoming County; NYSM 12080-12086. C. Data based on 30 specimens from the level of the Point Breeze Goniatite Bed of the lower Angola Shale (Locs. 72/6, 73/6, 74/6a, 75/6, 81/6, 82/6, and 83/6a) between Point Breeze, Lake Erie shore and Stony Creek, Wyoming County; material in NYSM. 147 148 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

have their discoidal form enhanced by lateral crushing, alto- Of Clarke’s cotypes, NYSM 3746 came from 45.7 m gether an unsatisfactory combination. In selecting NYSM (150 ft) “below the Portage sandstone, at the Lower Portage 3748 as lectotype, emphasis has been on the solid preserva- Falls on the Genesee River” and Clarke recorded that this tion indicating a rounded, rather than sharp, venter, and was the earliest record known to him. This locality (Genesee hence enabling comparison with the better quality material Gorge, Loc. 86, Wyoming and Livingston counties) is fig- described below. ured in Text-fig. 12 but the exact level of Clarke’s specimen Additional material.–The finest new material is a speci- is unknown; on our correlation (Text-fig. 3) the Table Rock men collected by J. Kralick (NYSM 16536) from the lower Sandstone in the Gardeau Shale at the top of the Lower Falls Angola Shale at Kennedy Gulf (Loc. 63/4). Ten specimens is equivalent to a level high in the Rhinestreet Shale of Erie collected in this investigation are included but only NYSM County, not to the Angola Shale as Clarke supposed. NYSM 12034 (as Sphaeromanticoceras aff. oxy, Text-fig. 44F) and 3747 from East Hill and NYSM 3748 (the lectotype desig- 12046-12047, 12133, 16569, and 16570 with confidence. natd here) from Caulkins Gully, Naples (Ontario County), Dimensions.–See Table 19. probably came from the Gardeau Shale and also NYSM Description.–The interpretation of this large species has E355, reported from the West Hill flags in the village of depended on the availability of material showing not only Naples referred to by Miller (1938: 95). the large discoidal outer whorls but also the earliest stages The new material from the Rhinestreet Shale is wholly (NYSM 16536) that are essentially those of Sphaeromantic- from the upper part of the unit. The collection from the Re- oceras rhynchostomum. This accords with the very large spec- lyea Creek Horizon (Loc. 60/3), and levels in the meter imen specimen figured by Clarke (and Miller, 1938: text-fig. below, at Relyea Creek (Wyoming County; Loc. 60), include 17) and the lectotype here designated. the large specimen NYSM 12034 (Sphaeromanticoceras aff. The umbilical width continues to be small with growth. oxy) and NYSM 16569 collected by L. V. Rickard. Most specimens show some degree of crushing, but the max- The best new material from the Angola Shale is NYSM imum whorl width continues to be near the shoulder and 16536 from the lower Angola Shale at Kennedy Gulf the flanks converge flatly to the venter. Clarke’s restored out- (Wyoming County; Loc. 63/4; Text-fig. 13) and from the line (NYSM 12305/4) shows a carinate venter, but the in- Point Breeze Goniatite Bed, NYSM 12046 and 3245/2, Big ternal molds of other specimens are often narrowly rounded Sister Creek (Loc. 73/5), NYSM 3229/2, 11, Cazenovia rather than acute, but this could reflect loss of the test; such Creek (Loc. 75/6), and NYSM 12047, the bed below the is the case in the largest specimen seen (NYSM 12034). Su- Point Breeze Goniatite Bed, Hampton Brook (Loc. 74/6a) tures of several of the larger specimens are illustrated here in Erie County. NYSM 12133 is from the eighth microcy- (Text-figs 44 C, E-F) and it will be noted that the ventrolat- clothem of the Angola Shale in Stony Creek (Loc. 83/8) in eral saddle can have a flater ventrad face (NYSM 12034: Wyoming County. Sphaeromanticoceras aff. oxy) or can be more symmetrical. Higher records in the upper West Falls Group include Several specimens show adult growth lines and these are Clarke’s 320-mm-diameter specimen from Stony Brook convex (NYSM 3747) with a very prominent ventrolateral Glen, Dansville (Steuben County; Loc. 87), mentioned by salient and what was formerly a latero-umbilical salient now Miller (1938: 95); this specimen could be from Angola Shale being only an undulation in the growth line course low on equivalents because we consider the Scraggy Bed level mark- the flanks (Text-fig. 44A). ing the base of the Angola to lie low in the section there. Discussion.–The source of the Sphaeromanticoceras oxy The Dansville area is also the source of NYSM 16570 from group is now clearly indicated as from S. rhynchostomum. the Nunda Sandstone or Wiscoy Sandstone equivalents in Because the range is similar, it raises the question of whether the roadcut I-390-Dansville (Steuben County; Loc. 87a). the former is the macroconch form and the latter a micro- Also reported from the Wiscoy Sandstone is Clarke’s giant conch and they represent a dimorphic pair. Further com- specimen (460-600 mm diameter) from Castile (Wyoming ment on this would require elucidation of the number of County; Miller, 1938: 95). The possibility that the Wiscoy whorls in each, and there is insufficient material available to Sandstone records could be the eastern representative of attempt this. Spheromanticoceras rickardi cannot be discounted. Distribution.–Middle Rhinestreet and Angola shales Range.–Frasnian: Playfordites and Neomanticoceras Geno- (Point Breeze Goniatite Bed) and equivalents (Gardeau zones UD I-I-J. Regional Zone of P. cf. tripartitus (22a) to Shale and West Hill Shale and Sandstone) (West Falls Sphaeromanticoceras rhynchostomum (22b). Conodont Zone Group) in Erie, Wyoming, Livingston, Steuben, and On- MN 11-?12. tario counties, and Nunda or Wiscoy sandstones (West Falls Group) in Wyoming, Livingston, and Steuben counties. HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 149

Table 20. Biometric data for Sphaeromanticoceras rickardi House & Kirchgasser, 1993, from the Hanover Shale.

Specimen D WW WH UW WW/WH UW/D

NYSM 12032 209 max. 140 60 (est.) 88 ca. 15 0.68 0.11 NYSM 12033 130 max. 92 47 52 — ca. 0.90 —

Sphaeromanticoceras rickardi House & Kirchgasser, 1993 neotype should be erected. He proposed for this a specimen Text-figs 48A-C; Table 20 from Belgium, rather than from the German type area.

Crickites holzapfeli (Wedekind, 1913). Matern, 1931b: text-figs 1a-b; House, 1978: 62. Sphaeromanticoceras rickardi House & Kirchgasser, 1993: 274, figs 5D-F.

Type material.–The species is based on IG 4449, a spec- imen from 560 m southeast of Romerée, chemin de Matagne-la-Petite, Belgium, and figured by Matern (1931b: text-figs 1a-b). New York material.–Only two specimens are known, a large, partly crushed, phragmocone (NYSM 12032) and a solid, incomplete specimen (NYSM 12033) showing septa and approximately half a whorl of body chamber, both from the 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 lines not seen. Suture with distinctive asymmetrical lateral saddle with nearly vertical dorsad face (LSh/LSw =1.17; Text-fig. 48C). Smaller specimen (NYSM 12033) showing half whorl of body chamber and adjacent proximal phragmocone showing involute form, deep umbilicus, and maximum Text-fig. 48. Sutures and cross sections of Sphaeromanticoceras, whorl width low on flanks that converge convexly to well- Crickites, and ?Archoceras. A-C. S. rickardi House & Kirchgasser, rounded venter. Complete suture not seen; growth lines in- 1993, from the Hanover Shale in New York and from Belgium. A. IG 4916, suture at 165 mm diameter based on a specimen from distinct. 2,700 m east-northeast of Roly (Sautour Sheet), Chemin de The specimen in the collection of G. Kloc (University of Viller's en Fagne, X 0.6. B. IG 4449, the "Neotype" figured by Rochester) with a lateral saddle with a less steep dorsad face Matern (1931b: text-fig. 1a-b) from 560 m south-southeast of to the lateral saddle was referred by us to Sphaeromanticoceras Romerée, Chemin de Matagne-la-Petite (Surice Sheet) at about aff. rickardi (see House & Kirchgasser, 1993: figs 5G-1. 100 mm diameter (suture for comparison), X 0.6. C. NYSM Discussion.–Growth lines are not shown on these speci- 12032, suture of a specimen at about 137 mm diameter from the mens so their earlier assignment to Crickites (Kirchgasser & Hanover Shale collected by L. V. Rickard at Glade Creek (Loc. House, 1981: 45, text-fig. 3) cannot be justified. Further, 91/7), Strykersville, Wyoming County, X 0.6. D-F. C. holzapfeli subsequent work on that genus has shown that the rotund Wedekind, 1913, whorl sections and a suture based on Wedekind's cotypes from Bicken, Germany (after Wedekind, 1913). Originals whorl sections of the C. holzapfeli and C. lindneri Group assumed to be natural size, X 0.6. G. ?Archoceras sp., NYSM continues through ontogeny to a large size. By contrast, 16587, cross section of a specimen from a concretion in the Sphaeromanticoceras rickardi becomes more compressed in Hanover Shale, Erie County, 4.7 m below the base of the Dunkirk later stages. In his revision of late Frasnian forms from Bel- Shale in Irish Gulf (Loc. 90). H. ?Crickites sp. juv., NYSM 16589, gium, Matern (1931b) illustrated type material. He argued from the same locality and horizon in Irish Gulf (Loc. 90). Scales that Wedekind’s type material of C. holzapfeli, being lost, a = 1 mm (G-H). 150 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

Table 21. Biometric data for Crickites lindneri (Glenister, 1958) from the Virgin Hills Formation (UWA), Western Australia, and the Hanover 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 0.20 MBC 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 0.19 ca. 39.6 28.1 20.9-15.2 — 1.34 — NYSM 16538 — 62.8 47.9- —— — —

Some of Wedekind’s type material however, appears to have Included species.–Genus restricted here to forms with survived, so this assignment is invalid. The suture of specified whorl outline: Crickites holzapfeli Wedekind (1913: Matern’s “neotype” (IG4449) from the Schistes de Matagne 72, pl. 7, figs 5-6, text-figs 14a-c); ?Manticoceras cordiforme (Text-fig. 48B), although not the same size as the New York Miller [1938: 82, pl. 21, figs 2-4; = ?M. septentrionale Miller, specimen, shows a similar shape, particularly in the upright, 1938: 102, pl. 23, figs 1-2, subjective), a more laterally com- asymmetrical lateral saddle at D = 100, LSh/LSw = 1.26. In pressed form and in the adult (fide House & Pedder, 1963, a rather larger specimen (IG4916) from near Roly (Text-fig. pl. 76, figs 10-11) with a very slight sinus on the lateral 48A), at D = 165, LSh/LSw = 1.24. The height of the me- salient]; M. lindneri Glenister [1958: 72, pl. 6, figs 1-2 dian saddle in this species is somewhat variable. (paratype), pl. 7, fig. 2, pl. 9, figs 1-2 (holotype), text-fig. 4, Distribution.–Type specimens, Europe, Matagne Shale 5B, C; one of the syntypes (UWA 35774) has a slight sinus of Belgium, and North America (New York), Hanover Shale on the lateral salient]; and M. neverovi Bogoslovsky (1958, (Wyoming County). pl. 3, fig. 1). Forms with very high lateral saddles, such as M. NYSM 12032, found by L. V. Rickard, from upper altaicum Bogoslovsky (1958: 100, pl. 4, fig. 1, text-fig. 25) Hanover Shale, Glade Creek, Strykersville (Wyoming should, perhaps, be separately placed. County; Loc. 91/7). NYSM 12033 was found loose just Discussion.–Matern (1931b) created a Belgian neotype above the base of the Hanover Shale also in Glade Creek for Crickites holzapfeli, but this is invalid because the original (Loc. 91). The G. Kloc specimen of Sphaeromanticoceras aff. types were then still available in Göttingen but they might rickardi came from 1.5 m above the base of the Hanover subsequently have been destroyed during World War II. It Shale in Walnut Creek (Cattatraugus County; Loc. 89a/5). seems best to base the species on the original figures of the Range.–Frasnian: NYSM 12032: Crickites Genozone type specimens from Adorf, Germany, figured by Wedekind UD I-L, Regional Zone of Sphaeromanticoceras rickardi (1913; refigured here, Text-figs 48D-F). (24b). Conodont Zone MN 12-?13. The type specimens The later whorls of some gephuroceratids show convex from Belgium from Archoceras Genozone UD I-K. Lower growth lines and if Wedekind’s subfamily is retained, it must Hanover specimens: NYSM 12033 and Kloc specimen of rest on the presence of convex growth lines from the earliest S. aff. rickardi: Crickites Genozone UD I-L, Regional Zone stages; these are known from approximately 9 mm diameter of C. lindneri (24a). Conodont Zone MN 12. in the type species (House, 1963: pl. 3, fig. a). Distribution.–Widely distributed. North America (west- Subfamily CRICKITINAE Wedekind, 1913 ern Canada, New York), Europe, Russia, North Africa, and Genus CRICKITES Wedekind, 1913 Western Australia. Type species.–Crickites holzapfeli Wedekind (1913: 72), Range.–Frasnian: Crickites Genozone UD I-L. Conodont by subsequent designation (Wedekind, 1917: 130). Zone MN 12-13. Diagnosis.–Subinvolute, medium- to large-sized manic- oceratids with convex growth lines from earliest stages with Crickites lindneri (Glenister, 1958) sinus on umbilical seam and broad salient over flanks. Text-fig. 49; Table 21 Whorl section distinctively rounded (rotund) to rounded- rectilinear with whorls converging convexly to rounded ven- Manticoceras lindneri Glenister, 1958: 72, pl. 6, figs 1-2, pl. 7, fig. ter (WW/WH = 1.0-1.3). Suture as in Manticoceras. 2, pl. 9, figs 1-2, text-figs 4, 5B-C. Sphaeromanticoceras lindneri. Becker, House & Kirchgasser, 1993: HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 151

Text-fig. 49. Sutures and cross sections of Crickites lindneri (Glenister, 1958) based on specimens from the lower Hanover Shale on Walnut Creek (Loc. 89a), Silver Creek, Chautauqua County. A-B. NYSM 16537, cross section and suture of a specimen collected by G. Klapper about 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. Klapper from 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. 152 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

Table 22. Biometric data for Probeloceras lutheri (Clarke, 1885) from the Cashaqua shale. See Text-fig. 35C for graph.

Specimen D WW WH UW WW/WH UW/D

NYSM 4067 ca. 29.6 — ca. 10.0 ca. 12.2 — 0.41 NYSM 13736 40.3 — 11.2 20.2 — 0.50 NYSM 12721 33.0 — 9.8 15.2 — 0.46 NYSM 12739 29.5 — 9.6 13.0 — 0.44 NYSM 12720 29.1 — 9.0 13.1 — 0.45 NYSM 12175 ca. 27.8 — 8.5 12.1 — 0.44 20.8 — 7.1 8.4 — 0.40 14.9 2.9 5.3 5.9 0.55 0.40 10.6 — 3.8 4.3 — 0.41 7.5 1.8 2.6 3.1 0.69 0.41 4.0 1.1 1.3 1.8 0.85 0.45 2.9 0.90 0.90 1.3 1.00 0.45 2.1 0.70 0.65 0.95 1.10 0.45 NYSM 12722 26.0 — 8.5 11.1 — 0.43 NYSM 12741 23.2 4.5 7.6 9.7 0.59 0.42 12.5 2.6 4.2 5.3 0.62 0.42 9.2 2.0 3.0 3.9 0.67 0.42 6.7 1.7 2.3 2.8 0.74 0.42 4.8 1.3 1.6 1.9 0.81 0.40 3.5 1.1 1.2 1.3 0.92 0.37 NYSM 12742 21.5 4.3 8.0 8.8 0.54 0.41 NYSM 12740 17.2 — 5.6 7.1 — 0.41 NYSM 3648 17.2 — 6.5 6.4 — 0.37 NYSM 12737 15.5 ca. 3.0 4.9 6.9 0.61 0.45 11.6 2.5 3.8 4.9 0.66 0.42 8.4 2.1 2.9 3.5 0.72 0.42 6.0 1.6 2.0 2.5 0.80 0.42 4.3 1.2 1.5 1.8 0.80 0.42 3.0 0.95 0.97 1.3 0.98 0.43 2.1 0.75 0.69 0.93 1.09 0.44 NYSM 12723 13.8 3.3 5.5 4.7 0.60 0.34 9.4 2.4 3.6 3.5 0.67 0.37 NYSM 12744 7.3 1.9 2.4 3.1 0.79 0.42 5.3 1.5 1.8 2.3 0.83 0.43

301-314. Dimensions.–See Table 21. Description.–NYSM 16537 showing cross section indi- Type material.–The type specimens of Glenister (holo- cating development of subinvolute specimen with rotund type, UWA 34765) are in the collections of the University whorl section from at least 2 cm diameter to maximum seen of Western Australia, Perth, where we have examined them. of over 10 cm (Text-fig. 49A). Suture typically gephurocer- New material.–Two loose specimens (NYSM 16537 and atid but with high, divided median saddle (Text-fig. 49B) 16538) collected by G. Klapper from the lower Hanover but rather symmetrical lateral saddle. NYSM 16538 (frag- Shale on Walnut Creek (Loc. 89a); one loose specimen (HM ment of larger specimen) showing suture with less high me- MBC 1943) collected by R. T. Becker from the same local- dian divided saddle and more asymmetrical lateral saddle ity; a fine specimen shown to us by G. Kloc from approxi- (Text-fig. 49C). Fragment of even larger specimen (HM mately Bed 89a/5 of the same section (Text-fig. 16). MBC 1943) showing subcircular cross section and growth

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 cross section 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 cross section 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, reversed suture 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 cross section, 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 based on a specimen from the Parrish Limestone, Conklin Gully (Loc. 47/2), near Naples, Ontario County, X 4. I. NYSM 12737 (continued right) HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 153

(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, Ontario County, X 4. K. NYSM 12742, cross section of a specimen from the Parrish Limestone, Griesa Hill Road (Loc. 46d), Naples, Ontario County, 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 shale capping the Parrish Limestone from the same locality and horizon, X 4. O, NYSM 12724, reversed suture at 24.8 mm diameter, from Cayuga Creek (Loc. 33/8), Cowlesville, Wyoming County, X 4. P,NYSM 12745, reversed suture at ca. 5.1 mm whorl height, from the Parrish Limestone, Snyder Gully (Loc. 46c/1), Woodville, Canandaigua Lake, Ontario County, X 4. Q, NYSM 3648, the holotype of Gephyroceras holzapfeli Clarke, 1898, suture at 16.8 mm diameter recorded as from the lower Portage shales, Eighteenmile Creek, near North Evans, Erie County, and probably from the Cashaqua Shale, X 5. 154 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

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) HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 155

lines of convex type with broad ventral sinus, arched lateral holzapfeli, the highest Frasnian goniatite zone-fossil in Eu- salient, and indications of umbilical sinus (Text-fig. 49E). rope, which also survived until the end-Frasnian extinction Discussion.–This species is characterized by the well- event. A thorough search of this level in Irish Gulf (Loc. 90) rounded whorl section throughout ontogeny as illustrated is required. That could enable the recognition of a terminal by Glenister (1958: 72, text-fig. 4; UWA 35766) although Regional Zone named after the species in New York. in that specimen, the outermost whorl (at 20.9 cm diameter) Distribution.–Upper Hanover Shale (West Falls Group), has become laterally compressed; it is comparable at similar Irish Gulf (Loc. 90/2), near North Boston (Erie County). diameters to the New York material (Table 21). As men- Bed 90/2 (Text-fig. 16) is a 2.0-2.5 cm-thick crinoidal and tioned above, one specimen showing growth lines in the gastropod-rich concretionary level, 1.76 m below the Frasn- Australian suite shows a slight sinus on the lateral salient ian-Famennian boundary and indicated by a concretion (UWA 35774), as does Crickites cordiforme, but the one symbol by Over (1997b: fig. 8). specimen from Walnut Creek (Loc. 89a) showing the Range.–Frasnian: Crickites Genozone UD I-L. Regional growth lines well (Text-fig. 49E) shows no evidence of this. Zone not assigned (24c). Conodont Zone MN 13. The specimens assigned to Manticoceras lindneri from the Cerro Gordo Member of the Lime Creek Formation of Family BELOCERATIDAE Hyatt, 1884 Iowa (Baker et al., 1986) lack the distinctively inflated whorl Diagnosis.–Gephuroceratoideans in which sutural ele- form of the Australian and New York Representatives of the ments become sigmoidal and usually angular with progres- species. These Iowa forms are part of the M. regulare fauna sive increase to seven adventitious and eight or more ventral of UD I-J (Becker & House, 2000) and could belong to umbilical lobes with many matched on dorsal suture. Rarely Sphaeromanticoceras. The M. regulare fauna is also found in with one ceratitic lobe. Shell laterally compressed and even- the Amana Beds in Iowa (Glenister et al., 2002). tually discoidal but with characteristic ventral band com- Distribution.–Virgin Hills Formation, Canning Basin, monly rather tabular. Growth lines biconvex. Western Australia. Lower Hanover Shale (West Falls Discussion.–The relations of this highly distinctive family Group), Walnut Creek (in the vinicity of Loc. 89a/4-5), at have now been clarified by the New York faunas, especially Silver Creek (Chautauqua County). the ancestral Probeloceras (see Kirchgasser, 1975), and doc- Range.–Frasnian: Crickites Genozone UD I-L. Regional umentation of the biostratigraphy of more complex forms Zone of C. lindneri (I-L1) (Western Australia: Becker et al., from the Canning Basin (Becker et al., 1993). This work has 1993; Becker & House, 1997), C. lindneri (24a) (New shown that the compilation of Yatskov (1990), which did York). Conodont Zone MN 13 (Australia), MN 12-?13 not have the advantage of the revision of non-Russian ma- (New York). terial nor a stratigraphy, is largely illusory. Nevertheless, we accept his new genus Naplesites Yatskov, 1990, as a conven- ?Crickites sp. juv. ient name for the intermediate group between Probeloceras Text-fig. 48H and Mesobeloceras Glenister, 1958, and hence to the most morphologically complex groups, Beloceras Hyatt, 1884, and New material.–NYSM 16589, single small specimen Ceratobeloceras House & Kirchgasser in House et al., 1985. showing a cross section but neither suture nor growth lines. Essential in the simplification has been the removal of eobe- Dimensions.–At D = ?4.4 mm, WW = 2.13 mm and loceratids thought by Yatskov to be contemporary but WH = 2.3 mm. shown to be late Givetian from work in Morocco (Becker & Description.–Poorly preserved cross section crushed in House, 2000a). earliest whorls. Subinvolute, final whorl section with There remains the question of the origins of the lineage. rounded venter and umbilical slopes with flat flanks con- The group seems clearly to have arisen from the Acanthocly- verging ventrally. Suture and growth lines not seen. meniidae, and a case can be made that Acanthoclymenia ge- Discussion.–This record, together with ?Archoceras sp. nundewa is the earliest, but the only known material is very (NYSM 16587), are the youngest Frasnian goniatites known small, and although some material of that taxon develops a in New York and thus have considerable importance. The sigmoidal ventrad face to the main lateral saddle (Kirch- rounded, stout cross section is reminiscent of Crickites 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 diameter and ca. 4.6 whorl height, Cazenovia Creek (Loc. 32/7), Spring Brook, Erie County, X 10. E. NYSM 12176, reversed suture of a flattened specimen 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. 156 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

Text-fig. 52. Sutures of Naplesites from New York State. A-E. N. iynx (Clarke, 1898), both probably from the lower Rhinestreet Shale at Naples, 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 holotype figured 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, X 6. H. NYSM 4066, a specimen from among Clarke's syntypes of Probeloceras lutheri (Clarke, 1885), reversed suture at 28 mm diameter, X 4.5.

the entry of the larger species, Probeloceras lutheri. moidal main lateral saddle, pointed lateral lobe (L), and sub- Distribution.–North America, Europe, North Africa, division of the dorsal suture giving a U1 lobe in adult. Shell Asia, and Western Australia. form laterally compressed with flattened venter. Growth Range.–Frasnian: Probeloceras to Crickites Genozones lines with strong forward projection across flanks. UD I-E-L (accepting P. lutheri as the base of the range). Distribution.–North America and Western Australia. Range.–Frasnian: Probeloceras Genozone UD I-E-F.Con- Genus PROBELOCERAS Clarke, 1898 odont Zones MN 5-6. Type species.–Goniatites lutheri Clarke (1885: 50), by monotypy. Probeloceras lutheri (Clarke, 1885) Diagnosis.–Beloceratids with no subdivision of ventral Pl. 24, Figs 5-9, Pl. 25, Fig. 1; Text-figs 35C, 50-51; Table 22 lobe beyond that found in acanthoclymeniids but with sig- Goniatites lutheri Clarke, 1885: 50, pl. 2, fig. 8. HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 157

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: figs 4C-D.

Type specimen.–Holotype, USNM 14993; NYSM 4067 (plastotype), figured here Pl. 24, Figs 7-8, from shale cap- Text-fig. 53. Sutures and growth lines of Tornoceras, from the Ge- ping Parrish Limestone, Cashaqua Shale, Conklin (Parrish) nundewa Limestone and Angola Shale, and Crassotornoceras, from Gully (Loc. 47/2), near Naples (Ontario County; Kirch- the Angola Shale in New York. A-B. T. uniangulare compressum gasser, 1975: 73). Clarke, 1897, based on a specimen from the Genundewa Lime- stone, Bethany Center (Loc. 24), Genesee County. A. NYSM Diagnosis.–Conch relatively large (to D = 60 mm), 12055, suture at ca. 22 mm diameter (estimated), X 1.3. B. NYSM highly evolute (subserpenticonic), widely umbilicate. Whorl 12056, growth line at ca. 28 mm diameter based on a specimen highly compressed (discoidal) with steep umbilical walls, from the same locality and horizon, X 1.3. C-D. T. cf. typum (G. subparallel, flattened flanks, weak ventrolateral furrows, and & F. Sandberger, 1851), based on a specimen from the Angola flat (tabular) venter that becomes rounded in outermost Shale in Cazenovia Creek (Loc. 75/6), Erie County. C. NYSM 12050, growth line at whorl height of 9.9 mm, X 2.7. D. Suture whorls. Suture (E1-Em-E1, L, U1, I) distinguished by highly asymmetrical, angular to acute lobes and saddles in chevron at a whorl height of 8.3 mm, X 2.7. E-F. C. aff. crassum (Matern, pattern. 1931), NYSM 12014, based on a specimen from the Point Breeze Dimensions.–See Table 22. Goniatite Bed, Angola Shale, Johnson Creek (Loc. 81/6), Wyoming County, X 3. E. Growth line. F. Course of constriction. Description.–Highly evolute and widely umbilicate through all growth stages. Whorls increasingly compressed beyond 2-3 mm diameter with flanks subparallel at nearly right angles to steep umbilical walls (Text-figs 35C, 50). across flanks and cross venter at right angles to revolving Weak ventrolateral furrows developing with whorl compres- keels. sion to latest stages. Venter distinctly flat or tabular by 1.5 Discussion.–The strongly asymmetrical and angular lobes mm diameter, becoming rounded beyond approximately 22 and saddles of Probeloceras lutheri serve to distinguish Probe- mm diameter. loceras from Acanthoclymenia and Prochorites. Prochorites is Suture across venter following pattern of Manticoceras distinguished by its concave venter. with ventrolateral lobes (E1) arising from bottom rather than Distribution.–Lower Cashaqua Shale (Sonyea Group), sides of ventral lobe (Text-fig. 50A; Kirchgasser, 1975: fig. widespread across outcrop belt between Erie and Steuben 4A). Sutural ontogeny marked by increasing angularity of counties. Unconfirmed records in Brallier Shale or equiva- lobes and saddles as characteristic chevron pattern develops. lents in Pennsylvania, West Virginia, and Virginia, and Ventral sides of lateral saddle becoming markedly crooked Woodmont Shale or equivalents in West Virginia and Mary- with distinct reflection toward umbilicus (Text-fig. 50). In land, and Millboro Shale of Virginia (Kirchgasser, 1975; latest stages, ventrolateral and lateral lobes becoming acute House 1978). Gogo Formation, Canning Basin, Western and lateral saddle nearly so. Australia (Becker et al., 1993, Becker & House, 1997). Growth lines of fine raised lirae with weak salient across Range.–Frasnian: Probeloceras to Prochorites Genozones umbilical shoulder, shallow broad sinus on lower third of UD I-E-F.Regional Zone of Probeloceras lutheri (19) to ?Pro- flank, strong, rounded salient on ventrolateral furrow, and chorites alveolatus (20) (New York). Regional Zones P.lutheri deep, tongue-shaped ventral sinus. Distinctive feature is pair lutheri (E1) to Sphaeromanticoceras affine (F3) (Canning of low, thread-like, revolving keels (“hyponomic tire” of Basin, Western Australia). Conodont Zones MN 5-6. Clarke, 1899a, b) inside the ventral margin (Kirchgasser, In New York, not known with certainty from upper part 1975: Pl. 2, Fig. 19). Ventral wrinkle layer consisting of uni- of Cashaqua Shale including the Shurtleff Septarian Hori- form, widely spaced parallel lines that sweep gently forward zon which yields Prochorites alveolatus. 158 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

Text-fig. 54. Sutures of Tornoceras spp. and sspp. from the Hamilton Group, Tully Limestone, and Genesee and Sonyea groups in New York and 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 the same 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 locality and horizon, X 3.5. D. T. cf. uniangulare, USNM 137721, suture at whorl height of 16.5 mm based on a specimen from the lower Windom Shale (Hamilton Group) at a quarry 1 mi (1.6 km) southeast of Lebanon Center, near Hamilton, Madison County, X 2. E. T. cf. uniangulare toward 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 15.3 mm 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, Livingston County, figured by Miller (1938: pl. 31, figs 5-6), X 2. H-J. T. arcuatum House, 1965, sutures of specimens from the Squaw Bay Limestone at Partridge Point, Alpena County, Michigan. H. USNM 96543a, reversed suture at 38.7 mm diameter, X 1.5. I. MMP 47509, suture at 27 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 estimated diameter of 41 mm, based on a specimen from the Platyceras Bed of the Tully Limestone, West Brook bed or member, (continued right) (Text-fig. 54 continued) fimbriata zone (Cooper & Williams, 1935), 0.5 mi (0.8 mm) northwest of Georgetown, Madison County, X 1. L. Tornoceras arcuatum, NYSM 12024, suture at ca. 24.5 mm diameter based on a specimen from the Crosby Sandstone in Sunset Point Gully (Py-16/1), East Branch, Keuka Lake, Yates County, X 1.5. M-O. P. concentricus, sutures of specimens from the Corell's Point Goniatite Bed of the Gowanda Shale at Corell's Point (Loc. 97), Brockton, Chautauqua County. M. USNM 137667, re- versed suture at 28 mm diameter, X 2.5. N. NYSM 11963, suture at 20 mm diameter, a composite drawing from adjacent septa, X 2.5. O. NYSM 11964, suture at 12.9 mm diameter, X 3. P. T. sp. USNM 96551D, suture at estimated whorl height of 29 mm based on a specimen from the Platyceras Bed of the Tully Limestone, West Brook bed or member, fimbriata zone (Cooper & Williams, 1935), 0.5 mi (0.8 km) northwest of Georgetown, Madison County, X 1.12. OS ICGSE:LT EOINGONIATITES DEVONIAN LATE KIRCHGASSER: & HOUSE

Text-fig. 55. Graphs showing the ontogeny of subspecies of Tornoceras uniangulare (Conrad, 1842) from New York; see Tables 23-25 for biometric data. A. T. u. uniangulare, data based on material listed herein and by House (1965), all from the Leicester ("Tully") Pyrite, Leicester, Livingstone County, and Canandaigua Lake. B. T. u. compressum Clarke, 1897, data based on material listed herein and by House (1965), all from the Genundewa Limestone; all the material above 12 mm diameter, and much of that below, is from Bethany Center (Loc. 24), Genesee County. C. T. u. obesum Clarke, 1897, data based on material listed herein and by House (1965), all from the Cashaqua Shale and most probably from the Shurtleff Septarian Horizon in the Honeoye Lake area. 159 160 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

Table 23. Biometric data for Tornoceras uniangulare uniangulare (Conrad, 1842) from the Leicester Pyrite. See Text-fig. 55A for graph.

Specimen D WW WH UW WW/WH UW/D

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 —

In the Canning Basin,Western Australia, Probeloceras 135, fig. 28B, pl. 12, fig. 11); House, 1962: 258. lutheri precedes Prochorites alveolatus as in New York, but Mesobeloceras naplesense. House & Kirchgasser, 1993: 276, 278, unlike in New York, it ranges upward and overlaps P.alveo- fig. 5J. latus in the form of P. aff. lutheri (with subacute venter) (F1), P.lutheri n. ssp. (in a sequence linking P.lutheri with Naple- Type material.–Yatskov (1990) based his interpretation on the holotype (NYSM 4072; formerly 12481/1), then the sites showing the gradual entry of an E2 lobe (F2) and P. cf. lutheri associated with Sphaeromaticoceras (Becker et al., only known specimen figured by Clarke (1899a, b: text-fig. 1993; Becker & House, 1997: 141; 2000: 128). 78, pl. 7, fig. 18) and Miller (1938: pl. 12, fig. 11, copied from Clarke). Reexamination of the holotype (by MRH in Genus NAPLESITES Yatskov, 1990 1959 and subsequently) shows that the suture on the lower Type species.–Probeloceras? naplesense Clarke (1898, flanks was incorrectly interpreted and that the suture there 1899a, b), by original designation (Yatskov, 1990: 43). actually shows a sharp L lobe and angular U2 probably ven- Diagnosis.–Beloceratids with single subdivision of ventral trad of the seam (revised here; Text-fig. 52G). Thus the as- saddle to form E lobe, and single subdivision low on flanks sigment of Miller to Neomanticoceras is incorrect as is the 2 diagnosis of Yatskov (1990). to form U2, and perhaps incipient E3 with up to four lobes between L lobe and seam; all lobes acute soon after develop- New material.–The suture of a better specimen found by ment; saddles subangular to acute. Intermediate in prolifer- WTK among the syntypes of Probeloceras lutheri, which is ation of acute lobes and saddles between Probeloceras and similar (NYSM 4066, formerly 12480/6), is illustrated here Mesobeloceras in a lineage that culminates in Beloceras Hyatt, (Text-fig. 52H). 1884. Dimensions.–Holotype: D = 61 mm; WH = 23.3 mm; Included species.–Naplesites anguisellatum (Chao, 1956), UW = 19 mm. N. housei (Montesinos & Henn, 1986), N. iynx (Clarke, Description.–The holotype is here illustrated photo- 1898, 1899a, b), N. cf. inyx (Becker et al., 1993), N. naple- graphically for the first time (Pl. 25, Fig. 6). The specimen sense (Clarke, 1898, 1899a, b), Naplesites sp. (Bogoslovsky et is crushed flat but consists of a thin, crushed pyritic mold of al., 1982: referenced by Becker & House, 2000: 129). the earlier whorls. The suture, illustrated by Clarke (1898, Distribution.–North America (New York; see Adden- 1899a, b: 105, text-fig. 78) and repeated by Miller (1938: dum), Europe, Asia, and Western Australia. text-fig. 28B), shows only a broadly rounded lateral saddle Range.–Frasnian: Mesobeloceras Genozone UD I-G. Sub- (L). It was noted by MRH (in 1959) that there was evidence division UD I-G1, range of Naplesites before the entry of of two angular lobes before the seam (Text-fig. 52G), and Mesobeloceras (UD I-G2) (Becker & House, 2000). this is confirmed in the new specimen (NYSM 4066; Text- fig. 52H). Naplesites naplesense (Clarke, 1898) Discussion.–Our reinterpretation of the lateral suture ren- Pl. 25, Fig. 6; Text-figs 52F-H ders the assignment to Neomanticoceras of Schindewolf (1936) and Miller (1938) untenable. A topotype of N. para- Anabeloceras pseustes Clarke, 1897: 53, nom. nud. et dubium. doxum, the type species of Neomanticoceras, from Probeloceras? Naplesense Clarke, 1898, 1899a, b: 105, text-fig. 78, Büdesheim, Germany, is figured here (Pl. 7, Figs 2-3) to il- pl. 7, fig. 18. lustrate differences. These New York specimens thus occupy Neomanticoceras naplesense. Schindewolf, 1936: 690; Miller, 1938: HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 161

an intermediate position between Probeloceras (but with an time. The inner whorls are pyritic and three-dimensional A2 and single ventral U lobe) and Naplesites iynx (with an but the outer whorls are crushed. The diagram of early su- acute A2 lobe and two ventral acute U lobes). Although tural ontogeny (Clarke, 1899a, b: text-fig. 77; Miller, 1938: Yatskov (1990) intended his genus Naplesites to apply to a text-fig. 29) is reproduced here (Text-figs 52A-E); in truth quite different form, we have now (with R. T. Becker) doc- the specimen on which this ontogeny is based (NYSM umented a succession in Western Australia through increas- 3583; Pl. 25, Fig. 4) could also represent early stages of ingly complex sutures to Mesobeloceras (with acute A2,A3, Naplesites naplesense, which would be indistinguishale from and even in some A4 lobes and with at least three ventral U N. iynx in early stages. lobes). Hence Naplesites has a restricted time span and the Discussion.–A rather similar and possible junior synonym name is biostratigraphically convenient. of Naplesites iynx is N. anguisellatum, the type species of Distribution.–Probably lower Rhinestreet Shale (West Chaoceras Yatskov, 1990, a genus here regarded as a junior Falls Group), in the area of Naples, Ontario, and Yates coun- synonym of Naplesites. ties. Reported from the “soft shales at Naples” and thought The sequence of morphologies between Probeloceras and to be from the lower Rhinestreet Shale. The reference to early Mesobeloceras in Western Australia, which include Naples in Clarke’s (1899c: 85) locality guide gives no help Naplesites cf. iynx and N. housei, indicates that the evolu- on the precise locality among sections in Parrish (Conklin) tionary stages were short lived (Becker et al., 1993; Becker Gully (Loc. 65a), Grimes Gully, or Tannery Creek. All have & House, 1997, 2000). The rarity of N. naplesense and N. been searched but no new material has been found. iynx suggest that they could have been closely contempo- Range.–Frasnian: Mesobeloceras Genozone (inferred) UD rary. I-G. Subdivision UD I-G1, range of Naplesites before entry Distribution.–Probably lower Rhinestreet Shale (West of Mesobeloceras (inferred). Regional Zone of N. iynx (21a). Falls Group), in the area of Naples, Ontario. and Yates coun- Conodont Zone unknown: MN ?6 or ?7. ties. Reported by Clarke from Portage shales at Naples but no locality or horizon details were given (see Distribution Naplesites iynx (Clarke, 1898) of Naplesites naplesense). Pl. 25, Figs 2-5; Text-figs 52A-E Range.–Frasnian: Mesobeloceras Genozone ((inferred) UD I-G. Subdivision UD I-GI, range of Naplesites before Beloceras iynx Clarke, 1897, nom. nud.; Clarke, 1898, 1899a, b: entry of Mesobeloceras (inferred). Regional Zone of N. iynx 103-104, text-figs 77-79, pl. 7, figs 11-16. (21a). Conodont Zone unknown: MN ?6, or ?7. Probeloceras iynx. Wedekind, 1918: 131. Eobeloceras iynx. Schindewolf, 1936: 690; Miller, 1938: 137-139, Order GONIATITINA Hyatt, 1884 text-fig. 29, pl. 30, figs 12-17. Naplesites clarkei Yatskov, 1990 [based on Clarke 1899a, b: pl. 7, Superfamily TORNOCERATOIDEA Arthaber, 1911 figs 15 (NYSM 3585) and 16 (NYSM 3586), text-fig. 78 This is the longest ranging superfamily of coiled ammonoids (NYSM 3583), and another specimen]. in the Devonian and, during its range, perhaps the com- Mesobeloceras iynx. House & Kirchgasser, 1993: 276, 278. monest. The superfamily is characterized by a generally sim- ple suture comprising a ventral lobe (E), an adult lateral lobe Type material.–Lectotype designated by Yatskov (1990: on the flanks, which develops adventitiously during on- 43) is NYSM 3581 (formerly 12060/1). Other cotypes togeny (A), a lobe centered on the seam (L), and a dorsal mentioned below, assigned elsewhere by Yatskov (1990) but lobe (D). Later forms can be more complex. Growth lines regarded here as synonyms of Naplesites iynx, include: are normally biconvex. Tornoceratids first appear in the late NYSM 3582 (Pl. 25, Fig. 2), 3583 (Pl. 25, Fig. 4), and 3584 Eifelian and continue to the latest Famennian, the last (not illustrated here), the umbilical portion of which is bro- records being from the Wocklumeria Stufe when extinction ken off, so the umbilical lobes are not seen, hence the as- is associated with the Hangenberg Event. The group is wide- signment is not clear. The original syntypes were formerly spread and cosmopolitan and is known from all continents numbered NYSM 12060/1-6. except for polar regions. Dimensions.–Lectotype, NYSM 3581: D = 25.4 mm, Three families are included: Tornoceratidae, Post- WW = 11.2 mm, UW = 7.3 mm. tornoceratidae, and Pseudoclymeniidae. Only the first is Diagnosis.–Distinguished from Naplesites naplesense by known in New York where it has a long record from beds additional sharp umbilical lobe on low flanks. immediately below the Cherry Valley Limestone (Hamilton Description.–Clarke (1899a, b) and Miller (1938) gave Group) to the Gowanda Shale (Canadaway Group). detailed descriptions of the type material and four of these specimens are here illustrated photographically for the first Family TORNOCERATIDAE Arthaber, 1911 162 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

Table 24. Biometric data for Tornoceras uniangulare compressum Clarke, 1897, from the Genundewa Limestone. See Text-fig. 55B for graph.

Specimen D WW WH UW WW/WH UW/D

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 —

Diagnosis.–Tornoceratids with imperforate, ornamented Subfamily TORNOCERATINAE Arthaber, 1911 ammonitella; umbilici and umbilicus open or closed with Tribe TORNOCERATINI Arthaber, 1911 shell form rotund and rounded to compressed and platy- Diagnosis.–Tornoceratidae with smooth shells, rounded conic or oxyconic. Adult shell usually smooth, rarely with venters, no ventrolateral furrows or tabular venters, and gen- falcate lirae, prominent ribs, nodes, or constrictions. Suture erally with umbilici. as described for superfamily but with simplification of lobes Discussion.–This group includes: Tornoceras Hyatt, 1984, in some (Tornia and Kirsoceras) and additional elements in the long-ranging nominal genus; Linguatornoceras House, others, a dorsal umbilical lobe (U), for example (in Loboto- 1965, a group with lingulate rather than asymmetric lateral rnoceras and Gundolficeras). (A) lobes; Epitornoceras Frech, 1902a, a late Givetian and Protoconch and ammonitella, where known, orna- early Frasnian oxycone; Crassotornoceras House & Price, mented with radial lirae; growth lines primarily biconvex 1985, a late Frasnian constricted form; and Lobotornoceras after nepionic constriction. Dorsal wrinkle layer spiralling Schindewolf, 1936, which adds a small saddle on the seam. backward from umbilicus to become almost rectiradiate over All of these occur in New York. Other genera are: Oxy- venter. tornoceras Becker, 1993a, a Famennian homeomorph of Epi- Discussion.–Several distinct groups are included within tornoceras; Domanikoceras Becker & House, 1993, a strange the family, which is thought to be derived from an unspec- early Frasnian form with convex growth lines from the ified Anarcestidae source. Only the family Tornoceratidae is Timan; and Semicheiloceras Becker, 1993a. These last three of concern here because the other families, Posttornocerati- genera are not known in North America. dae (developing additional ventral adventitious lobes) and Distribution.–Worldwide. Marcellus Formation (Hamil- Pseudoclymeniidae (with clymeniid-like lateral lobes), are ton Group) to Gowanda Shale (Canadaway Group) in New not represented in North America. York. Range.–Givetian MD II-A to Famennian UD VI-D. Subfamily PARODICERATINAE Petter, 1959 The Parodiceratinae forms the earliest group and comprises Genus TORNOCERAS Hyatt, 1884 Parodiceras, with only an incipient lateral lobe (A), as the Type species.–Goniatites uniangulare Conrad, 1842. earliest member, and Wedekindella and Trevoneites. The last Diagnosis.–Involute, well-rounded, or subglobular to two, reviewed by Becker & House (1994b), are only tenta- subplatyclonic shell with closed or nearly closed umbilicus; tively included here; both have well-developed lateral (A) without marked ventrolateral furrows. Adult suture with lobes but Wedekindella is a constricted form and Trevoneites EALD lobes only. Adult growth lines commonly biconvex has an open umbilicus. Only Parodiceras occurs in New York but biconvexity lost toward aperture in some species. in the Union Springs Shale (Werneroceras Bed) and Cherry Discussion.–Revision of the species of Tornoceras is still Valley Limestone (House, 1981b, 1978) of the Hamilton in progress. Several dozen names are taxonomically available Group, well below the levels dealt with in this monograph. for this genus and these will only be referred to in the follow- ing text if they are relevant to New York material. HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 163

Forms with small lingulate lateral lobes have been sepa- seo. This subspecies is only known with certainty from the rated as the subgenus Linguatornoceras, elevated herein to Leicester Pyrite. genus, as discussed below. The North American (House, Dimensions.–See House (1965), Table 23, and Text-fig. 1965), Russian (Bogoslovsky, 1971), and German (Becker, 52A. 1993a, 1995) species of Tornoceras have been subject to re- Description.–A description of this subspecies was given vision; the varied forms from elsewhere are mostly poorly by House (1965). known. A wide range of distinct sutural, growth line, and Discussion.–See House (1965) for detailed discussion of shell form types are included in the European faunas and ontogeny of this subspecies. there is no doubt that they will contribute to the discrimi- This group represents perhaps the commonest Devonian nation when restudied. Because the New York forms have goniatite. Comment is necessary, however, on the relative been relatively recently described (House, 1965), this ac- claims of Tornoceras uniangulare and Ammonites simplex (von count will be limited to new records and new statistical in- Buch, 1832: 42, pl. 2, fig. 8), the name commonly used for formation. this group in Europe. A general comparison between the Distribution.–Extraordinarily wide distribution. Known two species has long been claimed, and von Buch’s name for on all continents except Antarctica. In New York from Chit- the European form has a priority of ten years over that from tenango Shale of Marcellus Formation (Hamilton Group) New York. Von Buch’s specimen, however, has never been to Corell’s Point Goniatite Bed, Gowanda Shale (Canad- figured satisfactorily, and the small sutural thumbnail sketch away Group). given by him is quite inadequate to form a diagnosis. Fur- Range.–International range: Givetian MD II-A to Fa- thermore, von Buch recorded his specimen “am Rammels- mennian UD II-D. berge bei Goslar gefunden,” in which case the specimen is possibly a Foordites. Beyrich (1884: 212), however, asserted Tornoceras uniangulare uniangulare (Conrad, 1842) that this was an error, and that the specimen was from Pl. 26, Figs 1-4, 6; Text-figs 54A-C, 55A; Table 23 Büdesheim, although the grounds for this statement are not clear. If Beyrich was correct, then the specimen probably is Goniatites uniangularis Conrad, 1842: 268, pl. 16, fig. 4. a tornoceratid. But in the absence of a clear designation and Goniatites astarte Clarke, 1885: 29, pl. 2, figs 9-10. description, it seems that the species is critically undeter- mut. Loomis, 1903: 916-917, 919, Tornoceras uniangulare astarte. mined. It is true that Beyrich argued that the specimen fig- pl. 5, figs 1-2. Tornoceras (Tornoceras) uniangulare. Miller, 1938 (pars): 157-166, ured by the Sandberger brothers (1851: pl. 10, fig. 14) as pl. 31, figs 8-9, pl. 32, fig. 9, pl. 33, figs 5-6, pl. 35, figs 5-6. Goniatites retrorsus var. typus, was the same species, and it Tornoceras (Tornoceras) uniangulare uniangulare. House, 1965: has long been been interpreted as that in Germany, but there 104-106, text-figs 9A-H, 17E, pl. 7, figs 58-59, 61-63, 65-67. has been no formal designation of a neotype for A. simplex. It is notable that the species and subspecies referred to Type material.–The cotypes of Tornoceras uniangulare are Tornoceras uniangulare by House (1965), in which they show AMNH 5476 (two specimens). The one figured by House adult growth lines, give indication of a loss of biconvexity to- (1965: pl. 7, figs 65-67, and herein, Pl. 26, Figs 1-2), as ward the aperture; these are all Middle Devonian or early AMNH 5476/1, is designated lectotype if it is not the holo- Upper Devonian specimens and range to the Genundewa type (it is the only originally figured specimen). The original Limestone. On the other hand, T. simplex and T. typum are specimens of Goniatites astarte are NYSM 5661 and 5662. reported to have biconvex growth lines, the feature of early All of these specimens are from the Tully (now Leicester) stages of T. uniangulare. It could be that the later Tornoceras, Pyrite. from the Büdesheim level, differ in this respect from T. uni- New Material.–Although additional material of this va- angulare, but in the absence of modern work on European riety has been collected in the present survey, there is noth- Frasnian tornoceratids, this view is largely speculative. ing to add to the general description (House, 1965). Distribution.–Leicester Pyrite (Genesee Group), Leices- Additional statistical information is added to former data in ter (Livingston County). The two specimens of Hall the graphs (Text-fig. 55A). Sutures illustrative of the sub- (AMNH 5476) were from the Leicester (Tully Pyrite) near species are given in Text-figs 54A-C. The material of Loomis Leicester (Livingston County), but which of the several gul- (1903; NYSM 5661 and 5662) is from Canandaigua Lake. lies there (Loc. 17, 17a, or 18) is not recorded. The material Other material was listed by House (1965). Our collections of Loomis (NYSM 5661 and 5662) is from Canandaigua include NYSM 3180/1-3 from the Leicester Pyrite at Lake. Other material is as reported by House (1965). Fol- Taunton Gully (Loc. 17), Leicester, and 3181, 3187/1-3, lowing the work of Huddle (1974, 1981) and Baird & Brett and 3182 from the same level in Fall Brook (Loc. 19), Gene- (1986a), it is recognized that the Leicester Pyrite is a remanié 164 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

level that youngs progressively to the west and includes ma- Tornoceras cf. uniangulare (Conrad, 1842) terial derived from the upper Hamilton Group. toward Epitornoceras Range.–Probably Givetian MD II, now found derived in Text-fig. 54E Givetian MD III. There is a single specimen (NYSM 12049) referred here Tornoceras uniangulare (Conrad, 1842) Group from the Geneseo Shale, 2.4 m (11 ft) above the Leicester There is a range of poorly preserved material that does not Pyrite on Beards Creek, Leicester (Livingston County; Loc. allow the rigorous analysis applied to the various subspecies, 18/2). and that is referred to the Tornoceras uniangulare group in various forms of open nomenclature as listed below. The Tornoceras uniangulare compressum Clarke, 1897 precise determination of this material is impossible because Pl. 28, Figs 1-5; Text-figs 53A-B, 54F, 55B; Table 24 much of it is crushed and shows no sutures. Tornoceras uniangulare var. compressum Clarke, 1897: 54; 1898: 54; 1899a, b: 116-118, pl. 8, fig. 18 (pars); 1904: 358, 361, Tornoceras aff. uniangulare (Conrad, 1842) 371. Pl. 27, Figs 1-6 Tornoceras (Tornoceras) uniangulare (pars). Miller, 1938: 158, pl. 32, fig. 8. Material assigned to Tornoceras (T.) aff. uniangulare was de- Tornoceras (Tornoceras) uniangulare compressum. House, 1965: 109- scribed and figured by House (1965: pl. 10, figs 92-93, 97- 110, text-figs 11A-E, 18A. 100) from the upper meter or so of the Windom Shale (Moscow Formation, Hamilton Group) in Grove’s Creek Type material.–Clarke’s holotype (NYSM 4098) has been Quarry (Seneca County; Loc. 4; NYSM/CUPM 42402, described by House (1965). NYSM 12641-12643). There is a larger specimen of T. cf. New material.–Eighty specimens from the Genundewa uniangulare from the Windom Shale in a quarry 1 mi (1.6 Limestone at Bethany Center (Genesee County; Loc. 24), km) southeast of Lebanon Center, near Hamilton (Madison formed the basis of the description published earlier (House, County; USNM 137721) that might belong here. 1965). There are 47 specimens in the J. W. Wells collection (his loc. 101, NYSM/CU) and 33 specimens (including Tornoceras cf. uniangulare (Conrad, 1842) NYSM 12634-12640) collected by MRH and used in the Text-fig. 54D 1965 study. In addition, there are 20 specimens (including NYSM 12125-12127) more recently collected. From this The following material is referred to Tornoceras (T.) cf. uni- locality, approximately 30 additional specimens are now angulare: USNM 137721 from the Windom Shale (Moscow available. All are solid specimens, occasionally testate, pre- Formation, Hamilton Group) and near the top of the unit served in coarsely crystalline styliolinid limestone and many at Portland Point Quarry on the eastern side of Cayuga show sutures and growth lines. Lake, a large phragmocone and body chamber collected by Dimensions.–See House (1965) and Table 24. W. A. Oliver, Jr. (NYSM 12031). From the Geneseo Shale Discussion.–The holotype shows markedly biconvex (Genesee Group) on Beards Creek, Leicester (Livingston growth lines. The larger Bethany Center specimens differ in County; Loc. 18/1, 3108), over 30 poorly preserved showing a subdued ventrolateral salient and growth lines Tornoceras from 7.6 cm (3 in) below the first 2.5 cm (1 in) that are convex and almost rectilinear across the flanks (Text- black shale. From the Penn Yan Shale, specimens conform- figs 53A-B). The accompanying graph (Text-fig. 55B) plots ing with the species occur, including in the Lodi Limestone dimensions of the material available and adds further detail at Beards Creek (Loc. 18/4, 3153), a nodule bed 9.75 m (32 to that previously published. The criteria for the distinction ft) above the base of the Geneseo Shale. Other material of this subspecies remains as stated previously: “the consis- comes from a creek north of Abbey Gulf (Loc. 21), 2 mi tently high whorl-width proportion, the tendency to form a (3.2 km) northeast of Honeoye (Ontario County), from a tabular venter at high diameters, and the marked reduction level equivalent to Loc. 21/2 (Linden Horizon), 6.17 m in the prominence of the ventrolateral salient above 20 mm (20.25 ft) below the base of the Genundewa Limestone. diameter” (House, 1965: 110). Also conforming to the species is the following material Distribution.–Genundewa Limestone (Genesee Group; from the West River Shale in Livingston County: Beards Genesee, Livingston, Ontario, and Yates counties). Creek (Loc 18a/3), Leicester, 3128/1; Fall Brook (Loc. The holotype (NYSM 4098) is labeled “Genundewa 19a/2), Geneseo, 3117/2; and from higher in the same sec- Limestone, Canandaigua Lake.” The new material is mainly tion (Loc. 19a/4), 3118/2. from the embankment spoil extracted from the roadcut at HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 165

Table 25. Biometric data for Tornoceras uniangulare obesum Clarke, 1897, from the Cashaqua Shale. See Text-fig. 55 C for graph.

Specimen D WW WH-Wh UW WW/WH UW/D

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

Bethany Center (Genesee County; Loc. 24). So far, the sub- It is material of this subspecies that has provided the best species is only recognized in the Genundewa Limestone. No illustrations available of the wrinkle layer structures in attempt was made to search and collect in detail at other lo- Tornoceras (House, 1965: pl. 9, figs 78-79) and larger illus- calities. Specimens probably belonging to this subspecies trations of this are given here: general descriptions have been were collected by one of us (WTK) at the type locality of published by House (1971: 27). the Genundewa at Genundewa Point (Loc. 23b), Distribution.–Cashaqua Shale (Sonyea Group; Liv- Canandaigua Lake (Yates County; Loc. 23b/4, 3916; Loc. ingston and Ontario counties). 23b/5, 3915). The holotype (NYSM 4099) and also NYSM 11262- Range.–Frasnian: Koenenites Genozone UD I-B. Re- 11263, E 346 (approximately 200 specimens) are all bari- gional Zone of K. styliophilus kilfoylei (16). Conodont Zone tized and said to come from concretions in the Cashaqua MN 2. Shale in the Honeoye Lake area. It seems probable that they are from the Shurtleff Septarian Horizon (Text-fig. 10). Tornoceras uniangulare obesum Clarke, 1897 Other material is from the type locality of this horizon at Pl. 28, Figs 6-10; Text-figs 54G, 55C; Table 25 Shurtleff’s Gully (Loc. 41/1) (NYSM 16564, = D1365), near Livonia (Livingston County). Specimens more uncer- Tornoceras uniangulare var. obesum Clarke, 1897: 54; 1898, 1899a, tainly placed here include NYSM 11245 from Mount Mor- b: 116, pl. 8, fig. 17. ris (Livingston County), and NYSM 4097 from Naples (pars). Miller, 1938: 157-166, Tornoceras (Tornoceras) uniangulare (Ontario County). pl. 32, fig. 7. Tornoceras (Tornoceras) uniangulare obesum. House, 1965: 110- Range.–Frasnian: Prochorites Genozone UD I-F.Regional 112, pl. 8, fig. 72, pl. 9, figs 75-79, text-figs 12, 18B. Zone of P. alveolatus (20). Conodont Zone MN 6.

Type material.–The holotype (NYSM 4099) is a baritic Tornoceras arcuatum House, 1965 replacement from the Cashaqua Shale. A description was Pl. 26, Figs 7-10, Pl. 27, Fig. 16; Text-figs 54H-J, L given by House (1965). Tornoceras (Tornoceras) uniangulare (pars). Miller, 1938: 165, pl. New material.–Two hundred additional specimens from 33, figs. 3-4, pl. 34, figs 1-4; Stumm, 1951: 35. Clarke’s collection (NYSM E 346) of baritic material formed Tornoceras (Tornoceras) arcuatum House, 1965: 106-109, text-figs the basis for the account of this subspecies given earlier: also 10, 17D, pl. 8, figs 68-71. NYSM 11262-11263, 11245, 16564 (D1365), and NYSM 4097 and 4093. Type material.–The holotype (USNM 96543a) is from Dimensions:–See Table 25 and Text-fig. 55C. the Squaw Bay Limestone of Michigan as is all the material Description.–Little addition to the earlier account that formed the basis for the original description. (House, 1965) is required. Statistical data are plotted in New York material.–Two specimens (NYSM 12024 and Text-fig. 55C for comparison with the other subspecies. The 12051), both from the Crosby Sandstone at Keuka Lake regular, sharp fall in the WW/WH ratio has not been seen (Yates County), are assigned to the species. These are well- in other species of Tornoceras. Whereas there are patterns in preserved phragmocones but without evidence of shell struc- the WW/WH ratio that are discriminatory, it should be ture. noted that this material is all baritic shell replacement ma- Discussion.–The two New York specimens are illustrated terial, and hence the measurements refer to testate speci- here (Pl. 26, Figs 7, 10, Text-fig. 54L) so a detailed descrip- mens, whereas those for the subspecies uniangulare and tion is not warranted. The distinctively arcuate latero-umbil- compressum do not. The pattern of sutures is as in T. unian- ical saddle, which is nearly symmetrical, is diagnostic of the gulare. species and well shown in the new material. 166 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

House (1965: 108) described a specimen (USNM New York material: closed umbilicate with gentle slope 96551a) from the Platyceras Bed of the Tully Limestone, from umbilicus to very low shoulder; maximum width very West Brook bed or member, fimbriata zone (Cooper & low on flanks, which converge convexly and smoothly to Williams, 1935: 804-805), 0.5 mi (0.8 km) northwest of well-rounded venter. Suture showing gently arched latero- Georgetown (Madison County), as having a similar suture umbilical saddle with flat top (Text-fig. 53D). Growth lines form to this species. There are slight differences in Tornoceras bioconvex with subdued salient low on flanks and promi- cf. arcuatum, so far as the poor state of the specimen allows nent ventrolateral salient. Weak spiral ornament on flanks. comparison, in the different depth of the ventral lobe, and Discussion.–It is only the evidence of the late stage at the lesser symmetry of the latero-umbilical saddle (Text-fig. which biconvexity of the growth lines is retained that justi- 54K). Nevertheless, the T. arcuatum tornoceratid type could fies taxonomic separation from Tornoceras uniangulare, but well be a long ranging one, starting in the late Givetian insufficient material is available to demonstrate this with Pharciceras Stufe and ancestral to the Epitornoceras group, as complete satisfaction given that most specimens do not suggested earlier (House, 1965: 82), although the Michigan show growth lines. and Keuka Lake material is obviously later in age. Distribution.–Rhinestreet Shale and Angola Shale (West The occurrence of this species at levels with Koenenites in Falls Formation; Erie, Wyoming, and Livingston counties). the late Penn Yan Shale of New York suggests that the Rhinestreet Shale (approximate level of Relyea Creek Crosby Sandstone level of Torrey et al. (1932) is a close cor- Horizon): NYSM 12076, Cazenovia Creek (Erie County; relative of the Squaw Bay Limestone. Loc. 54/1); NYSM 12077, Stony Creek, Warsaw (Wyoming Distribution.—Squaw Bay Limesone, southern side of County; Loc. 61/1). Angola Shale (Point Breeze Goniatite Partridge Point, Alpena, Michigan. Crosby Sandstone, Penn Bed): NYSM 12050, Cazenovia Creek (Erie County; Loc. Yan Shale (Genesee Group; Yates County). NYSM 12024, 75/6). gully above Sunset Point (Loc. F, Py-16/1), Keuka Lake and NYSM 4093 (Pl. 27, Fig. 15), recorded from the NYSM 12051, Willow Grove Creek, Keuka Lake (Yates “Naples Formation” at Mount Morris (Livingston County), County; Loc. E, Py-14/1). is referred here with doubt. Range.–Frasnian: Koenenites Genozone UD I-B. Re- Range.–Frasnian: Playfordites to Neomanticoceras Geno- gional Zone of K. styliophilus styliophilus (16). Conodont zones UD I-I-J (also possibly earlier: Mesobeloceras UD I- Zone MN 2. G). Regional Zone of P. cf. tripartitus (22a) to Sphaeromanticoceras rhynchostomum (22b) [also possibly ear- Tornoceras cf. typum (G. & F. Sandberger, 1851) lier: Naplesites iynx (21a)]. Conodont Zone MN 11 to ?12, Pl. 27, Fig. 15; Text-figs 53C-D possibly earlier.

Goniatites retrorsus typus G. & F. Sandberger, 1851: pl. 10, figs 14, Genus LINGUATORNOCERAS House, 1965 14a-c. Type species.–Goniatites retrorsus var. lingua G. & F.Sand- berger, 1851. Type material.–The lectotype is the specimen figured by Diagnosis.–Species close to Tornoceras with well-rounded the Sandberger brothers and in the Wiesbaden Museum. to compressed-rounded form; distiguished by suture with New York material.–Three specimens from the small, deep, usually symmetrical, tongue-shaped lateral lobe. Rhinestreet and Angola Shales (NYSM 12050, 12076, and Included species.–Goniatites lingua G. & F. Sandberger 12077), preserved as internal molds in mudrock and crys- (1851: pl. 10, figs 20-21; 1852: 109); Tornoceras (Lingua- talline calcite and variously preserving part of the body tornoceras) aff. linguum House (1965: 112, pl. 9, figs 82, 93, chamber and phragmocone. Also a number of very poorly text-fig. 13E); T. clausum Glenister (1958: 92, pl. 15, figs 7- preserved specimens from the Angola Shale that might be- 9, text-figs 16a, c); T. guestphalicum Frech (1897: pl. 32a, long here but that cannot be dogmatically assigned. fig. 8, holotype noted by Becker, 1993a: 188, as refigured by Dimensions.–Lectotype: D = 45 mm, WW = 18.7 mm, House & Price, 1985); T. haugi Frech [1902b: 47, pl. 3(2), WH = 26 mm, Wh = 14.5 mm, UW = 0 mm. figs 20a-b]; T. pompeckji Wedekind [1918: 137, nom. nud., Description.–The Sandbergers’ drawing of the lectotype lectotype designated by Becker, 1993a: 189, as Frech, 1902: is accurate except that the lateral lobe is more asymmetrical pl. 4, figs 9b-c (as refigured by House & Price, 1985)]. and less tongue-shaped than indicated and the ventrad slope Discussion.–Bogoslovsky (1971: 52) did not favor sepa- of the latero-umbilical saddle is almost concentric. Growth ration of these forms even at the subgeneric level; this has, lines show a salient low on the flanks, so they are biconvex however, now been generally adopted. House (1965: 91) even at the large diameter of the specimen. thought that the group formed a discrete type of tornocer- HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 167

Table 26. Biometric data for Linguatornoceras aff. linguum (G. & F. Sandberger, 1851) from the upper Rhinestreet and lower Angola shales.

Specimen D WW WH-Wh UW WW/WH UW/D

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 ——

atid restricted to the mid-Frasnian to lower Famennian and imens in having flat, converging sides, rather than an ovoid was a useful time discriminator, and this has been confirmed whorl section, and in having a substantially deeper lingulate by Becker (1993a). There are other tornoceratids with very lateral lobe. It shares the latter character with L. clausum large lingulate lateral lobes that are nearly symmetrical; these from Western Australia. There are specimens from the seem too close to the Tornoceras s. s. stock to warrant taxo- holzapfeli Zone in Devon, England, figured here as L. aff. nomic separation. linguum (Text-figs 56E-F), which are younger in age and Distribution.–North America (New York), Europe (Eng- which have a flatter and broader ventrolateral saddle. Spec- land, France, Germany, Poland), Russia (Urals), North imens of L. linguum from Germany are illustrated here for Africa, and Western Australia. comparison (Pl. 27, Figs 13-14). Range.–Frasnian: Prochorites Genozone (UD I-F) to Distribution.–Rhinestreet Shale (Erie and Wyoming Paratorleyoceras Genozone (UD II-D). counties). Rhinestreet Shale (Relyea Creek Horizon or near equiv- Linguatornoceras aff. linguum (G. & F. Sandberger, 1851) alent): NYSM 12017 and 12021, Cazenovia Creek (Erie Pl. 27, Fig. 12, Pl. 33, Figs 1-2, 5-8; Text-fig. 56; Table 26 County; Loc. 54/1); NYSM 12018 and 12022, Relyea Creek (Wyoming County; Loc. 60/3); NYSM 12019, aff. Goniatites retrorsus var. lingua G. & F.Sandberger, 1851: pl. 10, Varysburg (Wyoming County; Loc. 57/2); NYSM 12020 figs 20-21; 1852: 109. and 12023, Johnson Creek (Wyoming County; Loc. 58/1). aff. House, 1965: 112, pl. Tornoceras (Linguatornoceras) linguum. Angola Shale (Point Breeze Goniatite Bed): possible rep- 9, figs 82-83, text-fig. 13E.

Type material.–There has been no further description of the type material published since it was originally figured by the Sandberger brothers. New York material:–Four specimens from the Rhinestreet Shale referred here (NYSM 12017-12020) and three others with some doubt (NYSM 12021-12023), all preserved as partial calcitic internal molds set in gray mudrock. Also NYSM 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- Text-fig. 56. Sutures and a growth line of aff. ing evenly to seam with no overhang. Suture as illustrated in Linguatornoceras lin- guum (G. & F. Sandberger, 1851) from New York and Devon, Text-figs 56A-B, D. Growth lines biconvex, with low umbil- England. A-C. Sutures and a growth line based on NYSM 12020 ico-lateral and ventrolateral salients, becoming more promi- from upper Rhinestreet Shale equivalents on Johnson's Creek (Loc. nent in outer whorls (Text-fig. 56C). 58/1), Wyoming County. A, Suture at ca. 20.2 mm diameter; B, Discussion.–These specimens seem identical with that suture at 13.8 mm diameter; C, growth line at ca. 14.8 mm diam- figured by Frech [1902a: pl. 3(2), figs 21a-b] from eter, X 1.5. D. NYSM 11248, suture at ca. 13 mm diameter based Büdesheim under the name Tornoceras simplex mut. ovata on a specimen collected by D. D. Luther at Relyea Creek (Gibson's (Münster, 1832) but, as Frech recognized, Münster’s name Glen) (Loc. 60), Wyoming County, X 2.7. E-F. Specimens from ovata refers to a different form altogether. Linguatornoceras the Saltern Cove Goniatite Bed (holzapfeli Zone), Saltern Cove, linguum, on the other hand, differs from the New York spec- Devon, England, sutures reversed for comparison. E. SM H 1526, X 3.7. F. BM c18461, X 3.7. 168 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

Table 27. Biometric data for Crassotornoceras aff. crassum (Matern, 1931) from the Angola Shale.

Specimen D WW WH UW WW/WH UW/D

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 ——

resentatives include NYSM 3230/13, Cazenovia Creek (Erie from levels probably corresponding to the former upper cor- County; Loc. 75/6). datum Zone (I gamma; now UD I-J), the source of the type NYSM 11248, collected by D. D. Luther in 1897, was species, Crassotornoceras ausavense crassum. The new material reported from Relyea Creek (Gibson’s Glen) (Wyoming from the Point Breeze Goniatite Bed of the Angola Shale is County; Locs 60 and 82), but whether it was from the upper probably of the same age. Crassotornoceras belgicum, from Rhinestreet or lower Angola is not known (House, 1965). the Assise de Matagne of Belgium, is from the holzapfeli Forms closely related to these specimens from New York Zone equivalents, perhaps UD I-K, hence later in age. occur in Europe in the upper cordatum Zone and holzapfeli There are also records of the genus in the lower Famenn- Zone of former usage, but they have not been adequately ian. Grüneberg (1925: 67, pl. 1, figs 11, 11a, 11b) referred studied. a specimen from the Cheiloceras Stufe of the Herzkamp Syn- Range.–Frasnian: In New York Playfordites Genozone cline to Steininger’s species Goniatites ausavensis Steiniger UD I-I to ?Neomanticoceras Genozone UD I-J. Regional (1853: 43, pl. 1, figs 9, 9a) but his figure seems closer to the Zone of P. cf. tripartitus (22a) to ?Sphaeromanticoceras rhyn- specimen figured by Frech (1902a: pl. 4, fig. 9b) from the chostomum (22b). Conodont Zone MN 11-?12. Cheiloceras Stufe at Nehden, Germany. Becker (1993a) has revised the lower Famennian occurences. Genus CRASSOTORNOCERAS House & Price, 1985 Type species.–Tornoceras ausavense crassum Matern, 1931a, Crassotornoceras aff. crassum (Matern, 1931a) by original designation. Pl. 33, Figs 9-20; Text-figs 53E-F; Table 27 Diagnosis.–Tornoceratids with globular inner whorls and very narrow umbilici with periodic constrictions following aff. Tornoceras ausavense crassum Matern, 1931a: 27, pl. 3, figs 14a- course of growth lines and numbering approximately four b. per whorl, increasing in some specimens in outer whorls to eight. Conch generally small. Sutures typically simple Type material.–The holotype of Crassotornoceras crassum tornoceratid; some with lingulate lateral lobe. was designated by Matern as the figured specimen (Senck. Included species.–Tornoceras ausavense crassum (Matern, Mus. XI 342a) from Büdesheim (UD I-J). The holotype is 1931a: 27, pl. 3, figs. 14a-b; type species); Crassotornoceras 7.6 mm in diameter with a whorl height in excess of width annissi House & Price (1985: 168-169, pl. 16, figs 12-13); and showing four constrictions. Because there is no evidence T. belgicum (Matern, 1931b: 9, text-fig. 2, refigured by of a small saddle centered on the seam, reference to Loboto- House & Price, 1985: text-fig. 3c, pl. 16, figs 1-3); ?C. iso- rnoceras ausavense is inappropriate. latum Becker (1993a: 186, text-fig. 68e, pl. 3, figs 11-12); New York material.–Seven specimens from the Angola C. nitidum Becker (1993a: 185, text-fig. 68f, pl. 3, figs 15- Shale (NYSM 12010-12015, 3229/17), none exceeding 17). 11.6 mm in diameter and all preserved as partial calcitic Remarks.–This distinctive group of small, constricted molds in gray mudrock. tornoceratids seems to characterize the upper part of the Dimensions.–See Table 27. Frasnian and the lowest Famennian. There are affinities Description.–Inner whorls at 3-5 mm diameter involute, shown by Crassotornoceras both with Linguatornoceras, Lo- globular, with well-rounded umbilical wall and subcircular botornoceras, and constricted species of Aulatornoceras, but whorl section. Whorl section becoming more compressed the exact relationships are obscure. with well-rounded umbilical wall and convexly flat flanks Distribution.–North America and Europe. that converge to well-rounded venter. Ornament with Range.–Frasnian UD I-J to Famennian UD II-C. This marked constrictions between 3 and 11 mm diameter, num- group is first known in the German Büdesheimer Schiefer bering four per whorl in early stages but reaching seven or eight in outer stages of body chamber. Constrictions follow- HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 169

ing course of growth lines (Text-figs 53E-F) and better Group 2. Epitornoceras. This has always been interpreted shown on internal molds than on outside of recrystallized as an oxyconic genus, but reillustration of the type species, tests where course marked only by shallow concavity, if at all. Epitornoceras mithracoides, by House (1978: pl. 10, figs 2-3; On internal molds, constrictions scarcely developed on um- Berl. Mus. c469) has shown that species not to be truly oxy- bilical wall or shoulder, deep across flanks, rather shallow conic, only laterally compressed with a tendency to become just dorsal of ventrolateral salient and again deeper in ventral subacute in larger whorls (shown by the second, unillus- sinus. Growth lines showing salient on lowest flanks, and in trated specimen in the Museum für Naturkunde, Berlin, early whorls a shallow sinus on mid-flanks, with lines pro- Berl. Mus. C470). Nor is the ventrolateral saddle as sharply jecting slightly to weak ventrolateral sinus and shallow ven- acute as shown in many illustrations. Nevertheless it seems tral sinus; in later whorls, lateral sinus relatively deeper and probable that E. mithracoides belongs to the early part of the ventrolateral sinus significantly more projecting, almost sub- stock that gives the true oxycones with sharp ventrolateral angular. saddles, such as E. peracutum, which occur in the late Givet- Sutures, seen only on smaller specimens (NYSM 12012), ian Pharciceras Stufe, and probably only in the lower parts. with umbilical lobe; generally convexly and backwardly di- Included are: E. mithracoides (Frech, 1887: 30, pl. 2, fig. 1; rected ventral suture with weak asymmetrical lateral saddle. House, 1978: pl. 10, figs 2-3); E. peracutum (Hall, 1879: Ventral lobe small, passing convexly to rounded ventrolateral pl. 69, fig. 8, pl. 74, fig. 13: holotype, NYSM 4091, figured saddle. by House, 1965: text-fig. 13C); E. aff. peracutum (figured by Discussion.–This form is very close to Crassotornoceras House, 1978: 60, pl. 6, figs 7-8, 11-12, text-fig. 7B); E. aff. crassum, but the constrictions in that species continue over peracutum (figured by Harris, 1899: pl. 6, fig. 35; refigured the umbilical shoulder and, even at a larger size, are less fre- by House, 1965: text-fig. 13H, pl. 8, fig. 74, NYSM/CU quent. One of the paratypes of Aulatornoceras loeschmanni 39652). Frech [1902a: 49, pl. 4 (5), fig. 9a; Berlin Mus. C 475/1] is Group 3. Oxycones are again derived from the conser- more similar, but this specimen shows constrictions passing vative Tornoceras stock in the early Nehdenian. These have over the umbilical shoulder also and, as with C. crassum, been named Oxytornoceras (Becker, 1993a) with a type more lingulate sutures of a type not apparent in the smaller species, by original designation, of T. acutum (Frech, 1902a: specimens from New York that show sutures. 47, pl. 3(2), fig. 17). A sequence of forms giving a clear link Distribution.–Point Breeze Goniatite Bed (or approxi- from typical Tornoceras to oxyconic forms with an acute ven- mate equivalent), Angola Shale (Erie and Wyoming coun- ter has been elucidated by the late Dr. H. Makowski in col- ties). NYSM 12010 and 12011, Hampton Brook (Erie lections from the early Famennian of the Holy Cross County; Loc. 74/6a); NYSM 12012-12013 and 3229/17, Mountains of Poland. These might be quite homeomorphic Cazenovia Creek (Erie County; Loc. 75/6); NYSM 12014, with the true Epitornoceras in that the ventrolateral saddle Johnson Creek (Wyoming County; Loc. 81/6); NYSM can, unusually, become acute. 12015, Relyea Creek (Wyoming County; Loc. 82/6). Material of Group 2 is only available for description Range.–Frasnian: Neomanticoceras Genozone UD I-J. from New York. The range and distribution data given Regional Zone of Sphaeromanticoceras rhynchostomum (22b). below apply only to Group 2. Conodont Zone MN 11-?12. Discussion.–New York specimens agree in the oxyconic form with the usual interpretation of the genus, but it Genus EPITORNOCERAS Frech, 1902a should be emphasized that the comparison with the type Type species.–Goniatites mithracoides Frech, 1887. species is based on the similar large, lingulate symmetrical Diagnosis.–Tornoceratids with oxyconic form in adult lateral lobes and not on whorl form. The earliest of the or, in some, earlier stages; with or without acute ventrolateral known New York representatives (NYSM/CU 42401) still saddle. Otherwise like Tornoceras. shows a marked asymmetry in the lateral lobe (Text-fig. Included species.–There are three quite different oxyconic 57F). The holotype of E. peracutum (Text-fig. 57A; NYSM stocks derived from “normal” Tornoceras of the T. uniangu- 4091) and the other New York and West Virginia specimens lare/simplex stock. The groups are: are all closely united in sutural form, but there is variation Group 1. Gen. nov. Small oxycones with Tornoceras uni- in depth of lateral lobe. New York is the only place in the angulare-style sutures, but with open umbilicus and world where there is evidence bearing on the time-succession “pinched” and acute ventral margin. Seen only from a spec- of the group. NYSM/CU 40105, from the Geneseo Shale at imen in the collection of G. Kloc (University of Rochester) Hubbard Quarry (Seneca County; Loc. 27b), is the largest from the Wanakah Shale (Hamilton Group), northwest of known tornoceratid from North America, if not the world. Bethany Center (Genesee County). The original figures of Oxytornoceras acutum show the 170 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

boundary. The specimen from the Harrell Shale of West Vir- ginia (House, 1978) with ?Pharciceras galeatum Wedekind, 1917, probably correlates to the earlier part of the New York range.

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.

Text-fig. 57. Sutures, cross section, and outline of Epitornoceras Type material.–Frech’s original material is in the Museum spp. from the Genesee Group in New York, the Harrell Shale in für Naturkunde (Berl. Mus. c469, 470) and one specimen West Virginia, and Oberscheld, Germany. A. E. peracutum (Hall, has been refigured by House (1978). 1879), NYSM 4091, suture of the holotype at 41 mm whorl New York material.–One large crushed specimen from height, Ithaca Shale and Sandstone, Ithaca, Tompkins County, X the Geneseo Shale (NYSM/CU 42401) preserved in dark 0.5. B-C. E. aff. peracutum. B. USNM 186052, reversed suture of gray shale. a specimen figured by House (1978: 34) from the Harrell Shale, Dimensions.–NYSM/CU 42401: D = ca. 200 mm; WH near Landes Post Office, West Virginia (see Weary & Harris, 1994), X 0.5. C. NYSM/CU 39652, suture of a specimen figured = 120 mm; Wh = 64 mm; UW = 0 mm. by Harris (1899: pl. 6, fig. 35) at ca. 80 mm diameter from the Description.–The body chamber seen to 340°; shell Ithaca Shale and Sandstone, Ithaca, Tompkins County, X 0.5. D- crushed; shell form not discernable. Involute with closed E. E. cf. peracutum. D. NYSM/CU 369, cross section based on a umbilicus. Suture as shown in Text-fig. 57F. Aperture not solid specimen from the upper Geneseo Shale or lower Penn Yan seen but evidence suggesting salient low on flanks; another Shale, southern side of Fir Tree Point (Loc. 25), Seneca Lake, inferred in usual ventrolateral position. southeast of Rock Stream, western side of Seneca Lake, Yates Discussion.–This specimen is compared to Epitornoceras County, X 0.63. E. NYSM/CU 40105, outline (drawn by J. W. mithracoides because of the large size and the similar, slightly Wells) showing the apertural margin of a specimen reaching ca. asymmetric large lingulate lateral lobe that distinguishes it 240 mm diameter from the upper Geneseo Shale at Hubbard from the later form, E. peracutum. Epitornoceras mithracoides Quarry (Loc. 27b), Seneca County, X 0.13. F. E. cf. mithracoides (Frech, 1887), NYSM/CU 42401, suture of a specimen from 12.2 seems to have its closest analog among the mainstream m (40 ft) below the top of the Geneseo Shale in Lodi Glen (Loc. tornoceratids, with Tornoceras arcuatum, the type material 27), Seneca County, X 0.5. G. E. mithracoides, BM c469, reversed of which is from the much later Squaw Bay Limestone of suture of one of the cotypes of Frech (1887a) from Grube Eibach, Michigan. Rather similar forms, recorded as T. cf. arcuatum, Oberscheld, Germany, X 0.5. are already known in the Tully Limestone (House, 1965, and herein), but these have narrower lateral lobes and more highly arched and symmetrical latero-umbilical saddles than growth lines well and they differ from the large New York E. mithracoides. Presumably something like T. cf. arcuatum specimen described here (Text-fig. 57E) in not having a gave rise to Epitornoceras as indicated previously (House, well-developed salient on the lower flanks. This could be 1965: text-fig. 1, a figure that was not intended to imply one means of distinguishing the early Famennian group that the Michigan T. arcuatum belonged to the Tully level). from the true Epitornoceras. Distribution.–Upper Geneseo Shale (Genesee Group), Distribution.–North America (New York and West Vir- Lodi Glen (Seneca County; Loc. 27). ginia) and Europe (Germany). NYSM/CU 42401 was collected by J. W. Wells in Lodi Range.–Givetian MD II to Frasnian UD I-A. The early Glen (Loc. 27; his loc. 111g), 12.2 m (40 ft) below the top Frasnian oxyconic tornoceratids correctly referred to Epi- of the Geneseo Shale and approximately 24 to 27.5 m (80- tornoceras occur in Germany only in the Pharciceras Stufe 90 ft) above the Tully Formation. (Frech, 1902a, b). In New York, forms assigned to the genus Range.–Givetian MD III (?C-?D). Regional Zone of Epi- appear in the upper part of the Geneseo Shale (Genesee tornoceras peracutum (15a), but occurring before the zonal Group) and records continue to at least the middle Penn species. Yan Shale, thus they range above the Givetian-Frasnian HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 171

Table 28. Biometric data for Epitornoceras peracutum (Hall, 1876) from the Geneseo Shale and Ithaca Shale and Sandstone.

Specimen D WW WH-Wh UW WW/WH UW/D

NYSM 4091 ——41- —0 — — (holotype) CU 40105 ca. 240 — ca.142- ca. 74 0—— NYSM 369 75 18 45- —0 0.40 —

Epitornoceras peracutum (Hall, 1876) Group NYSM/CU 39652 is only given as Ithaca in Tompkins Pl. 34, Fig. 8; Text-fig. 57A; Table 28 County, and it can only be presumed that they are from lower levels of the Ithaca Formation. The rare specimens of Goniatites peracutus Hall, 1876: pl. 69, fig. 8, pl. 74, fig. 13; 1879: Epitornoceras known elsewhere from above the Geneseo 463, pl. 69, fig. 8, pl. 74, fig. 13. Shale are from well up in the Penn Yan Shale [3846/1, Clarke, 1898, 1899a, b: 118, text-fig. 96. Tornoceras peracutum. Taunton Gully (Loc. 17/4); 3853/4, Fall Brook-Dewey Hill Tornoceras (Epitornoceras) peracutum. Frech, 1913: 19. Tornoceras (Tornoceras) peracutum. Miller, 1938: 155-156, pl. 31, (Livingston County; Loc. 19/5)]. These Penn Yan Shale figs 1-2. specimens from Livingston County are poorly preserved. Epitornoceras peracutum. House, 1965: 118-119, text-fig. 13c, pl. USNM 186052, Epitornoceras aff. peracutum, figured by 8, figs 73-74. House (1978: 60, text-fig. 7B, pl. 6, figs 7-8, 11-12; suture refigured here, Text-fig. 57B), is from the Harrell Shale, 0.8 Type material.–The holotype (NYSM 4091) has been de- km (0.5 mi) south of Landes Post Office (Grant County), scribed by Miller (1938) and House (1965). West Virginia. New material.–Other specimens from the Geneseo Shale Range.–Givetian MD III (?C-?D) to Frasnian UD I-A. conforming to the species include: NYSM/CU 40105 (Text- Regional Zone of Epitornoceras peracutum (15a) to Ponticeras fig. 57E), an extremely large, crushed specimen, and perlatum (15b). NYSM/CU 369, a solid specimen (Text-fig. 57D, Pl. 34, Figs. 6-7), both referred to Epitornoceras cf. peracutum, and Genus LOBOTORNOCERAS Schindewolf, 1936 less well preserved material and specimens seen in the field Type species.–Goniatites ausavense Steininger, 1853, by but uncollectible. NYSM/CU 39652, the specimen figured original designation (Schindewolf, 1936: 689), a form from by Harris (1899: pl. 6, fig. 35), is refigured here (Text-fig. Büdesheim, Germany; topotype figured by Schindewolf 57C, Pl. 34, Fig. 5) and is referred to E. aff. peracutum. (1936) and neotype of House (1978), SM H9932, herein Dimensions.–See Table 28. Text-figs 58C, F. Discussion.–In view of the illustrations given here, much Diagnosis.–Tornoceratids with closed or nearly closed additional comment is unnecessary. The cross section of umbilicus, well-rounded compressed to subplatyconic form, NYSM/CU 396 (Text-fig. 57D) shows the progressive onset rarely with ventrolateral furrows; some with prominent lat- of oxyconic form that is achieved after approximately 25 eral constrictions. Suture as in Tornoceras but with additional mm diameter. There is some variation in the depth of the saddle centered on umbilical seam (EAL’UD). Growth lines lateral lobe, and NYSM/CU 39652 (Text-fig. 57C) is distin- biconvex. guished from the holotype by the lesser depth of the lateral Included species.–A neotype for Lobotornoceras ausavense lobe. (Steininger, 1853: 40, pl. 1, fig. 11) was selected and figured Distribution.–Upper Geneseo Shale to middle Penn Yan by House (1978: pl. 9, figs 8, 14; Sedgw. Mus. H 9932). Shale and equivalents in the Ithaca Formation (Genesee Additionally there is L. hassoni House (1978: 59, pl. 8, figs Group; Tompkins County), westward to Livingston County. 6-7, 10-11, text-figs 11E-G; holotype USNM 239891, Harrell Shale (Grant County), West Virginia. paratypes 239892-239896, 240505) and L. aff. hassoni de- NYSM/CU 40105, the very large specimen from at or scribed herein. Famennian species formerly grouped here near the top of the Geneseo Shale at Hubbard Quarry have been assigned to the genera Falcitornoceras and Exo- (Seneca County; Loc. 27b). tornoceras by Becker (1993a). NYSM/CU 369, from upper Geneseo Shale or lower Discussion.–This genus appears to arise from Tornoceras Penn Yan Shale reported from “between Rock Stream and s. s. by the addition of a small saddle centered on the um- Rock Landing,” southern side of Fir Tree Point, Seneca Lake bilical seam. This happened at least twice but the Frasnian (Yates County; Loc. 25). species show a distinctive, broad, subsymmetrical lateral The source of the holotype (NYSM 4091) and lobe, whereas later Famennian homeomorphs show more 172 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

Table 29. Biometric data for Lobotornoceras aff. hassoni House, 1978, from the West River Shale.

Specimen D WW WH UW WW/WH UW/D

NYSM 12029 7.4 — 4.8 ? 0.3 — 0.04 NYSM 12030 6.7 max. —— — — — NYSM 12030 — 2.1 3.24 ? 0.3 0.65 —

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 recorded from a styliolinid limestone, possibly close to the level of the Genundewa Limestone, in the Harrell Shale in West Vir- ginia (House, 1978). Lobotornoceras aff. hassoni is described herein 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 the Northwest Territories of Canada (House & Pedder, 1963). Text-fig. 58. Sutures and cross section of Lobotornoceras spp. from Lobotornoceras aff. hassoni House, 1978 New York, West Virginia, Northwest Territories, and Germany. A- Pl. 34, Figs 1-4; Text-figs 58A-B; Table 29 B. L. aff. hassoni House, 1978. A. NYSM 12029, reversed suture based on a specimen from the upper West River Shale on Seneca aff. Lobotornoceras hassoni House, 1978: 59, pl. 8, figs 6-7, 10-11; Point Creek (Loc. 23a/3), X 4.6. B. NYSM 12030, suture and text-figs 11E-G. whorl section of a specimen from the upper West River Shale on Whetstone Brook (Loc. 26a/1), near Honeoye, Ontario County; Type material.–The holotype of Lobotornoceras hassoni suture at 3.2 mm whorl height and cross section at 3.4 mm whorl (USNM 239891; Text-fig. 58E) and paratypes (USNM height, X 4.6. C. L. ausavense (Steininger, 1853), suture of a spec- imen from Büdesheim, Germany, figured by Schindewolf (1936), 239892-239896, 240505) are from a styliolinid-rich nodule reversed for comparison; magnification not stated. D. Falci- in the Harrell Shale, from 4.1 mi (6.6 km) south of Landes tornoceras aff. bilobatum (Wedekind, 1908), GSC 16955, suture at Post Office (Grant County), West Virginia. 34 mm diameter figured by House & Pedder (1963: 529) from New York material.–Two specimens only from the West near Carlson's Lake, Northwest Territories, X 0.8. E. L. hassoni, River Shale in Ontario County: NYSM 12029, a crushed USNM 239891, suture of the holotype at 5.8 mm whorl height phragmocone preserved as an incomplete pyritic internal from the Harrell Shale south of Landes Post Office, West Virginia mold, from Seneca Point (Loc. 23a/3); NYSM 12030, a (see Weary & Harris, 1994, for locality details), X 4. F. L. solid, incomplete, pyritized internal mold, from Whetstone ausavense, SM H 9932, suture of the neotype from the Büde- Brook (Loc. 26a/1). sheimer Schiefer at 210 m south-southeast of Büdesheim Church, Dimensions.–See Table 29. Germany, X 4.6. Description.–The larger specimen (NYSM 12029) shows approximately 18 sutures in the last half whorl. Shell form Discussion.–Sutural form of the larger of these specimens difficult to determine due to crushing but the maximum approaches the holotype of Lobotornoceras hassoni but both whorl width appears to lie approximately one third of the specimens differ in the absence of evidence of a well- distance from the umbilicus to the venter; the flanks slope rounded form and flattish venter. Both specimens differ evenly to the umbilicus without any shoulder; the venter ap- from L. ausavense in showing no constrictions, and from Fa- pears to be abruptly rounded and could have a flattening. mennian homeomorphs in the form of the suture. Suture as illustrated in Text-fig. 58A. The smaller specimen Distribution.–Middle to upper West River Shale (Gene- (NYSM 12030; Text-fig. 58B) indicates a shell form again see Group; Ontario County). with the maximum whorl width approximately one third of Range.–Frasnian: Timanites Genozone UD I-C. Regional the distance from the umbilicus to the venter with no um- Zone of Koenenites beckeri (17b). Conodont Zone MN 4. bilical shoulder; toward the venter the flanks converge con- vexly and there is a well-rounded ventrolateral shoulder and Tribe FALCITORNOCERATINI Becker, 1993a a flatly rounded venter; suture as illustrated in Text-fig. 58B. Diagnosis.–Tornoceratidae similar to the Tornoceratini HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 173

Table 30. Biometric data for Phoenixites concentricus (House, 1965) from the Gowanda Shale. See Text-fig. 59A for graph.

Specimen D WW WH-Wh UW WW/WH UW/D

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

but commonly with ventrolateral furrows in earliest stages fig. 14, pl. 9, figs 84, 86-88, pl. 10, figs 101-110, pl. 11, fig. and falcate early ornament, and with lateral constrictions. 138. Some with additional umbilical saddle. Phoenixites concentricus. Becker, 1993a: 198. Discussion.–In this group, Becker (1993a, 1995) in- cluded Phoenixites, Falcitornoceras House & Price, 1985 Type material.–The holotype [NYSM 11963 (6679D)] [type species Goniatites (Tornoceras) subundulatus var. falcata and nine other specimens from the Gowanda Shale were de- Frech, 1887], and Gundolficeras Becker, 1995 (type species scribed by House (1965). Lobotornoceras bicaniculatum Petter, 1959), the last two New material.–Eight additional specimens are available commonly developing a saddle on the umbilical seam. (listed below). Here we follow the taxonomy of Becker (1993a, 1995), Dimensions:–See Table 30 and Text-fig. 59A. but with some misgiving because detailed ontogenies have Discussion.–The new material adds extra statistical data not been described for many of the included forms. Most that are plotted in Text-fig. 59A for comparison with other of the many species of Tornoceras have early stages that are known tornoceratids. Other details remain as given in the quite unknown and their assignment is put in question primary description in which details of the earliest stages when a genus (Phoenixites) is erected based only on charac- were given. The diagnostic suture of the species, with its ex- ters of the early stages. ceedingly steep ventrad face to the latero-umbilical saddle Distribution.–Worldwide. In New York, represented by and the sharp turn over the ventral crest of that saddle in Phoenixites concentricus in the Gowanda Shale (Canadaway adult stages, is illustrated in Text-figs 54M-N. There is no Group). new evidence regarding sexual dimorphism speculated upon Range.–Probable worldwide range: Givetian MD II to earlier (House, 1965: 115). Too little is known of the early Famennian UD VI. In New York: Famennian: UD II-C. stages of most Tornoceratidae for this species to be separated from the Tornoceratini with any confidence. Genus PHOENIXITES Becker, 1993 Distribution.–Corell’s Point Goniatite Bed, Gowanda Type species.–Tornoceras frechi Wedekind, 1918, by orig- Shale (Canadaway Group; Chatauqua, Cattaraugus, and inal designation (Becker, 1993a: 198). Wyoming counties). Diagnosis.–Like Tornoceras but with ventrolateral grooves Only known from the Corell’s Point Goniatite Bed of in the early stages and some with constrictions. the Gowanda Shale. Material listed by House (1968) and Included species.–See Becker (1993a, 1995). NYSM/CU 3292/1, 6, 14, Corell’s Point on Lake Erie Distribution.–North America, Europe, Russia, and Shore (Chatauqua County; Loc. 97); NYSM 12052, Big In- North Africa. dian Creek (Cattaraugus County; Loc. 101); NYSM 12053 Range.–Upper Givetian MD III-A to Famennian UD II- and 12054, and NYSM/CU 3278/20, Cattaraugus Creek C. (Cattaraugus County; Loc. 104); NYSM 16565 and 16566 (Pl. 29, Figs 11-12), Java (Wyoming County; Loc. 109). Phoenixites concentricus (House, 1965) Poorer material of this species has been noted at almost every Pl. 29, Figs 1-15; Text-figs 54M-O, 59A; Table 30 outcrop of the Corell’s Point Goniatite Bed inland of Lake Erie. Tornoceras (Tornoceras) concentricum House, 1965: 113-115, text- Range.–Famennian: Cheiloceras (Cheiloceras) Genozone 174 B LEISO MRCNPLOTLG o 374 no. PALEONTOLOGY AMERICAN OF ULLETINS

Text-fig. 59. Graphs showing ontogeny of Phoenixites, Truyolsoceras, and Cheiloceras, all based on material from the Corell's Point Goniatite Bed, Gowanda Shale, in western New York. See Tables 30 and 34-35 for biometric data. A. P. concentricus (House, 1965), as Tornoceras concentricum, based on material listed herein and by House (1965) from Corell's Point (Loc. 97), Big Indian Creek (Loc. 101), and Cattaraugus Creek (Loc. 104), Chautauqua and Cattaraugus counties. B. T. bicostatum (Hall, 1843), as Aulatornoceras biocostatum, based on material listed herein and by House (1962) from Corell's Point (Loc. 97) and Walnut Creek (Loc. 99b), Chautauqua County. C. C. (Cheiloceras) amblylobum (G. & F. Sandberger, 1851), based on material listed herein and by House (1962) from Corell's Point (Loc. 97), Chautauqua County. HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 175

Text-fig. 60. Sutures and cross sections of Truyolsoceras and Aulatornoceras spp. from the Gowanda Shale and Angola and Hanover Shales in western 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), Erie County, X 4. E. A. rhysum (Clarke, 1898), NYSM 4092, suture and growth line of the lectotype at 5 mm diameter, based on a specimen said to come from the Hanover Shale at Java, Wyoming County, X 9.2. F. A. paucistriatum, NYSM 12094, cross section of same specimen as 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.

UD II-C. Regional Zone of C. amblylobum (25). but until European species are more precisely defined, both taxonomically and stratigraphically, little progress will be Tribe AULATORMOCERATINI Becker, 1993a made in the elucidation of the evolution of the group. It Genus AULATORNOCERAS Schindewolf, 1922 seems possible to recognize three groups of aulatornoceratids Type species.–Goniatites auris Quenstedt, 1846, by origi- as follows: nal designation (Schindewolf, 1922: 188). Neotype chosen Group A. These include the type species and are charac- by House & Price (1985: pl. 17, figs 1-4; SM H9942) from terized by a closed or nearly closed umbilicus and ornament Büdesheim. without very strong ribs or constrictions but with adults that Diagnosis.–Tornoceratidae with ventrolateral furrows, are rather smooth-sided although the growth lines can be commonly flat-sided with tabular venters and closed or prominent on the flanks and commonly form festoons on nearly closed umbilicus in early stages to subinvolute in the venter. Included species: Aulatornoceras auris (Quenst- adult. Ornament forming strong ribs, or with bunching of edt, 1846: 64, pl. 3, figs 7a-c); ?A. auris bickensis (Wedekind, prominent growth lines or radial bands; in some with 1918: 137; Matern, 1931a: 31); A. eifliense (Steininger, growth-line festoons on venter increasing in frequency in 1849: 27; 1853: 43, pl. 1, figs 3, 3a, but not 2, 2a); A. ei- adult; most with periodic constrictions. fliense posterior Becker (1993a: 216, text-figs 77e-f, 78d, pl. Discussion.–Miller (1938: 142) drew attention to the two 9, figs 7-10); A. sandbergeri (Foord & Crick, 1897: 112 pro groups of morphologies then included within Aula- G. & F.Sandberger, 1851: pl. 10, figs 17, 17a; not Goniatites tornoceras. He distinguished between a group with open and undulatus Brown, 1841); and A. lepiferum Becker (1993a: a group with closed umbilici. There is clearly a wide range 215, text-figs 77c, 78b, pl. 8, figs 11-13, pl. 9, fig. 1). of very different forms included within the genus at present, Group B. These are forms with markedly open umbilici 176 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

in early whorls, and open umbilici in later whorls, and that show the development of ventral festoons (Pl. 31, Fig. 3). are extremely heavily ribbed in the early whorls although None of the specimens shows the suture. ribbing declines in prominence on the outer whorls. In- Discussion.–This is a “waste-basket” group used for large cluded species: Aulatornoceras paucistriatum (d’Archiac & de aulatornoceratids with nearly closed umbilici (when seen) Verneuil, 1842: 339, pl. 25, figs 8, 8a, 8b); A. rhysum and united by the development of festoon-like grooves pe- (Clarke, 1898: 121, text-fig. 100, pl. 8, fig. 14; NYSM riodically on the venter that follow the growth line course. 4092); and A. serriense Becker (1993a: 213, text-figs 77a-b, This is a feature of large specimens generally referred to 78a, pl. 8, figs 14-18). Aulatornoceras auris from Büdesheim. The possibility that Group C. These are closed umbilicate, rather smooth- outer whorls of A. paucistriatum, A. rhysum, and A. eifleinse surfaced aulatornoceratids with periodic constrictions. In- are included here cannot be discounted, although the last cluded species: Aulatornoceras constrictum (Steininger, 1853: would be unlikely. The one cross section (Text-fig. 60G) 43, pl. 1, fig. 9); A. auriforme Oppenheimer (1916: 174, pl. suggests a different early stage than these species. From the 1, fig. 2; see Becker, 1993a); A. keyserlingi Müller (1956: 49 Gowanda Shale, Truyolsoceras bicostatum does not appear to pro Keyserling, 1846: 277, pl. 12, fig. 3; not Goniatites cinc- develop festoons and T. clarkei is more widely umbilicate. tum Münster, 1843); and A. loeschmanni (Frech, 1902a: 49, The alternative to using the open nomenclature followed pl. 4 (5), figs 9a-c) here would be to erect new names, but this would be inap- Discussion.–The first two groups are well represented in propriate at present. Taxonomic revision and detailed de- the New York State Devonian and the third group not at all. scription of the European species, particularly from Some use is made of European names in the account that Germany, are needed. follows because there can be little doubt as to the close rela- Distribution.–Angola Shale and Hanover Shale (Erie and tionship of the forms concerned, but this is bedeviled by the Wyoming counties). fact that, despite wide use of the names in Germany, few of Type material from Büdesheim, Gemany. This species the German type specimens have been figured photograph- name has been used widely in Germany and internationally ically since they were originally described. for Frasnian aulatornoceratids. The exact source of the ma- Distribution.–North America, Europe, Russia, North terial of Quenstedt from Büdesheim is not known, but is Africa, and Western Australia. probably from the main locality southeast of the church, Range.–Frasnian: UD I-I to Famennian UD II-D. and hence a level in the late cordatum Zone is most probable, but the species could well range higher. Aulatornoceras auris (Quenstedt, 1846) Group Becker et al. (2000: 78, fig. 4) reported Aulatornoceras Pl. 31, Figs 2-3; Text-fig. 60G cf. auris from the Lyaiol Formation in Timan, Russia (UD I-I). Goniatites auris Quenstedt, 1846: 64, pl. 3, figs 7a-c. In New York, the records are from the Angola Shale (below the Point Breeze Goniatite Bed) and lower Hanover Type material:–The type material of this species has not Shale, but not also from upper Rhinestreet Shale as stated by been redescribed or figured since its first description. The House & Kirchgasser (1993: 276, 279, fig. 7) and Becker & specimen figured by Quenstedt was said to come from House (2000: 129). Angola Shale: NYSM 12113, Hampton Büdesheim, Germany, and a large number of horizons there Brook (Erie County; Loc. 74/3); NYSM 12114 and 12115, yield aulatornoceratids. Despite the fact that the species is Sheldon Creek (Erie and Wyoming counties; Loc. 78/4b). the type-species of the genus, it is not critically determinable Hanover Shale: NYSM 12117 and 12118, Beaver Meadow by modern standards. It is used here in a wide sense. Creek, Java (Wyoming County; Loc. 92/3). New York material.–Generally poorly preserved large Range.–In New York: Frasnian: Neomanticoceras Geno- specimens are included here (NYSM 12113-12118) from zone (UD I-J) to Archoceras Genozone (UD I-K). Regional the Angola and Hanover Shales, none of which are com- Zone of Sphaeromanticoceras rhynchostomum (22b), Delph- plete. Most show evidence of the body chamber only, all are iceras cataphractum (23). Conodont Zone MN 11-12. preserved in mudrock with some shell replacement by cal- cite, and some have crystalline calcite infilling. Aulatornoceras eifliense (Steininger, 1849) Description.–This material does not warrant detailed de- Pl. 31, Figs 7-13, 16-20; Table 31 scription, but illustrations are given (Pl. 31, Figs 2-3, Text- fig. 60G). One solid specimen (NYSM 12113) shows the Goniatites eifliensis Steininger, 1849: 27; 1853: 43, pl. 1, figs 3, proportions of the inner and outer whorls (Text-fig. 60G); 3a. other specimens are less well preserved. Most specimens Goniatites retrorsus var. undulatus G. & F. Sandberger, 1851: pl. 10, fig. 9. HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 177

Table 31. Biometric data for Aulatornoceras eifliense (Steininger, 1849) from the Cashaqua Shale.

Specimen D WW WH UW WW/WH UW/D

NYSM 12107 13.5 8.0 — 0—— NYSM 12106 7.9 ca. 4.7 —0 —— NYSM 12105 5.1 2.4 —0 — —

Tornoceras eifliense. Wedekind, 1918: 137. by Becker (1993a) except that it is further restricted to spec- Aulatornoceras eifliense. House & Kirchgasser, 1993: 276. imens showing the same evidence of a laterally compressed Aulatornoceras eifliense eifliense. Becker, 1993a: pl. 9, figs 4-6. form as indicated by Steininger’s figure. Distribution.–Cashaqua Shale (Sonyea Group; Erie, Type material.–Steininger’s material has neither been re- Genesee, and Livingston counties). described nor rediscovered. Wedekind (1917) selected one Specimens include: NYSM 12105 and 12111, Smoke of the specimens figured by Steininger (1853: pl. 1, fig. 3) Creek (Erie County; Loc. 31/5); NYSM 12106, Cazenovia and this could be construed as selection of a lectotype. But Creek (Erie County; Loc. 32/9); NYSM 12110, Murder this has been overtaken by the selection as a neotype by Creek (Genesee County; Loc. 34/6); NYSM 12109, Buck Becker (1993a: 370) of a specimen in the Wiesbaden Mu- Run Creek (Livingston County; Loc. 39/17); NYSM seum figured by the Sandberger brothers (1851: pl. 10, fig. 12107, Shurtleff Septarian Horizon, Shurtleff’s Gully (Liv- 19) from Büdesheim. ingston County; Loc. 41/1). New York material.–Seven specimens from the Cashaqua A poorly preserved specimen (NYSM 12112) from the Shale (NYSM 12105-12111) including some earliest stages Angola Shale, Lake Erie Shore (Farnham Creek; Loc. 71/9a), preserved in pyrite but larger specimens preserved as baritic shows a similarly closed umbilicus and unornamented form replacements of the shell. but cannot be placed here with any confidence; it is referred Dimensions.–See Table 31. to Aulatornoceras aff. eifliense. Description.–Inner whorls with closed umbilicus from Range.–In New York: Frasnian: Probeloceras Genozone approximately second whorl. Whorl form at first showing (UD I-E) to Prochorites Genozone (UD I-F). Regional Zone significant lateral compression, with convex flanks, ventro- of Probeloceras lutheri (19) to Prochorites alveolatus (20). lateral furrows, and convex ventral band conforming to total Conodont Zone MN 5-6. whorl outline (NYSM 12105; Pl. 31, Figs 10-11). Whorl width increasing relatively and by largest diameter seen Aulatornoceras aff. eifliense (Steininger, 1849) (NYSM 12107; Pl. 31, Figs 18-20), with whorl form dif- Pl. 31, Figs 4-6, Pl. 32, Fig. 14; Table 32 fering from Büdesheim types in being less compressed and more inflated. Sutures showing V-shaped ventral lobe and aff. Goniatites eifliensis Steininger, 1849: 27; 1853: 43, pl. 1, figs asymmetric lateral lobe in early whorls at approximately 5 3, 3a. mm diameter (NYSM 12105; Pl. 31, Fig. 11); suture not aff. Aulatornoceras eifliense eifliense. Becker, 1993a: pl. 9, figs 4-6. seen at larger diameters. Shell surface generally smooth, without constrictions, ribs, or festoons on venter. Ornament New York material.–Five specimens from the Rhinestreet consisting of growth lines alone, with striae forming lateral and Angola Shales. NYSM 12102-12104 preserved as solid sinus, passing forward to form lappet centered in ventrolat- phragmocones in pyrite and mudrock, and NYSM 12116 in eral groove, and passing back to U-shaped ventral sinus. mudrock. Discussion.–Wedekind used the species name Tornoceras Dimensions.–See Table 32. eifliense for specimens like Aulatornoceras auris but with a Description.–Small subglobular aulatornoceratids with closed umbilicus and without constrictions. That is the sense closed umbilicus, inflated whorl form, and weak ventrolat- in which it is used here and conforms to the neotype selected eral furrows. Growth lines (NYSM 12104) with salient cen-

Table 32. Biometric data for Aulatornoceras aff. eifliense (Steininger, 1849) from the Angola Shale.

Specimen D WW WH UW WW/WH UW/D

NYSM 12103 7.1 4.4 —0 —— NYSM 12104 5.9 3.8 — ca. 0 —— 178 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

tered on rounded umbilical shoulder, with broad lateral Dimensions.–See Table 33. sinus, lappet in ventrolateral groove, and deep U-shaped Description.–Small, openly umbilicate aulatornoceratids ventral sinus. Sutures (NYSM 12103) with ventral V-shaped with strongly evolute inner whorls (NYSM 12096, see Pl. lobe, linguiform lateral lobe, and flat-topped latero-umbil- 32, Figs 7, 9; NYSM 12094, see Text-fig. 60F) and de- ical saddle. NYSM 12116 (Pl. 31, Fig. 4) showing distinc- pressed whorl section; changing to subevolute outer whorls tively paired growth lines on flanks. with less depressed section and deep umbilicus at largest di- Discussion.–These are discriminated from Aulatornoceras ameters known (9.1 mm, NYSM 12094). Ornament char- eifliense on the basis of the more rotund form; they are also acterized by deep, incised grooves periodically on flanks (Pl. later in age. 32, Figs 5-6) that sweep concavely forward to a slight ven- Distribution.–Upper Rhinestreet and lower Angola Shale trolateral groove. On flanks, these grooves or groove-like (West Falls Group; Erie and Wyoming counties). constrictions increasing from approximately eight per whorl Rhinestreet Shale: NYSM 12116, Kennedy Gulf at 4 mm diameter to ten or more at 7 mm diameter, forming (Wyoming County; Loc. 63/1). Angola Shale: NYSM part of finer ribbing pattern at larger diameters when ribs 12103 and 12112, Lake Erie Shore (Farnham Creek) (Erie can exceed 16-18 per whorl. In middle whorls, festoon-like County; Loc. 71/9a); NYSM 12102, Hampton Brook (Erie grooves appearing along growth-line sinus of venter, but County; Loc. 74/6a); NYSM 12104, Sheldon Creek only weakly developed, if at all, along ventrolateral groove; (Wyoming County; Loc. 78/4b). some appearing as periodic spiral grooves along ventrolateral Range.–In New York: Frasnian: ?Playfordites Genozone line (NYSM 12100). Growth lines appearing to follow (UD I-I) to Neomanticoceras Genozone (UD I-J). Regional course of lateral grooves, sweeping forward to slight ventro- Zone of ?P. cf. tripartitus (?22a) to Sphaeromanticoceras rhyn- lateral furrow, and back to U-shaped ventral sinus; low chostomum (22b). ?Conodont Zone MN 11 and ?earlier. salient formed low on flanks. Suture showing narrowly V- shaped ventral lobe, deep, lingulate lateral lobe, and latero- Aulatornoceras paucistriatum umbilical saddle that is distinctly flat in its course to a lobe (d’Archiac & de Verneuil, 1842) centered on seam (Text-fig. 60D). Pl. 32, Figs 2-13; Text-figs 60D, F; Table 33 Discussion.–It could be that some of the material de- scribed by Clarke under the name Tornoceras bicostatum be- Goniatites paucistriatum d’Archiac & de Verneuil, 1842: 339, pl. longs here [Clarke, 1899a, b: pl. 8, figs 4-8; NYSM 25, figs 8, 8a, 8b. 4081-4085, refigured by Miller, 1938: pl. 30, figs 5-9, as T. (Aulatornoceras) bicostatum]. This material, recorded as being Type material.-The original material has not been re- from near Angola and probably from the lower Angola described, and it was not located during a search of de Shale, is excluded from the account given above, which is Verneuil’s collection in the l’Ecole des Mines in Paris made based on well-localized material. by one of us (MRH) in 1958. The original material was Attention has been drawn earlier to the possibility that recorded as from Adorf and Oberscheld, Germany. outer whorls included here under Aulatornoceras auris could New York material.-Several specimens from the Angola belong to this species but this cannot be determined unless Shale (including NYSM 12094-12101 and BMNS E inner whorls are present. It is clear, however, that the early 22464), mostly preserved in concretionary mudrock with whorl of specimens referred to A. auris at Büdesheim are dif- some crystalline calcite filling of internal whorls. One spec- ferent. imen is a siliceous shell replacement (NYSM 12096). One specimen now included here (BMNS E 22464) was Diagnosis.–This name is used here for openly umbilicate formerly assigned to Aulatornoceras rhysum; the specimen Aulatornoceras conforming to the type illustrations with was said to be from the Hanover Shale at Hampton Brook strong ribs or grooves on the flanks, especially in the inner (Erie County), but in view of the material collected of this whorls. species 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.

Specimen D WW WH UW WW/WH UW/D

NYSM 12094 9.1 5.1 4.8 2.6 1.06 0.29 NYSM 12095 7.5 ca. 4.4 3.3 2.6 1.33 0.35 NYSM 12096 4.2 ca. 2.1 ca. 1.18 1.8 1.78 0.43 HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 179

the horizon was the lower Angola Shale. Genus TRUYOLSOCERAS Montesinos, 1987 The horizon of the original German type material is not Type species.–Tornoceras sandbergeri Foord & Crick, known exactly; the localities of Adorf and Oberscheld pro- 1897, nom. nov. pro Gon. undulatus G. & F. Sandberger duce aulatornoceratids at several levels in the Frasnian (1851: 101, pl. 10, figs 17, 17a), non Brown, 1841. The (Adorfian). Most records of the species elsewhere have not Sandbergers’ specimen is from the Frasnian of Büdesheim, included descriptions thorough enough to enable compari- Germany. son with the New York material. Diagnosis.–Tornoceratids with nearly closed umbilicus; Distribution.–Lower Angola Shale (West Falls Group; laterally compressed adults with ventrolateral furrows, sim- Erie and Wyoming counties). ilar to Aulatornoceras but without the festoons or constric- All New York specimens, except three, from either the tions. Point Breeze Goniatite Bed or concretionary bed immedi- Included species.–Truyolsoceras sandbergeri (Foord & ately below it. BMNS E 22464, from an uncertain horizon Crick, 1897); T. bicostatum (Hall, 1843: 246, text-fig. in Hampton Brook as noted above; NYSM 12100, Hamp- 107(8); holotype AMNH 5888/1); and T. clarkei (Miller, ton Brook (Erie County; Loc. 74/3); 3226/3, Sheldon Creek 1938: 80-81, pl. 14, figs 15-17; cotypes NYSM 5652- (Erie and Wyoming counties; Loc. 78/4b). 5654). Point Breeze Goniatite Bed or bed immediately below: Discussion.–Separation of these forms from Aula- NYSM 12095-12099, 12101, and 3235/4, 9, 13, Hampton tornoceras without considerably more work on biostratigra- Brook (Erie County; Loc. 74/6a); 3228/1, 4, 5, 9, 3229/15, phy and evolution of the aulatornoceratids was premature. 16, 25, 34, 35, 38, and 3230/4, 7, 8, 10, Cazenovia Creek As it stands, Truyolsoceras is used for aulatornoceratids with- (Erie CountyLoc. 75/6). out the morphological extreme festoons of the auris Group, Range.–In New York: Frasnian: Neomanticoceras Geno- ribbing of the paucistriatum Group, or constrictions as in zone UD I-J. Regional Zone of Sphaeromanticoceras rhyn- the constrictum Group (see groups discussed under Aula- chostomum (22b). Conodont Zone MN 11. tornoceras above). Because such forms occur in the Givetian it could be the stock from which others are derived. Aulatornoceras rhysum (Clarke, 1898) Pl. 32, Fig. 1; Text-fig. 60E Truyolsoceras bicostatum (Hall, 1843) Pl. 27, Figs 7-10, Pl. 30, Figs 2, 4-14; Tornoceras rhysum Clarke, 1898, 1899a, b: 121, text-fig. 100, pl. Text-figs 59B, 60A-C, H; Table 34 8, fig. 14. Tornoceras (Tornoceras) rhysum. Miller, 1938: 156-157, pl. 30, fig. Goniatites bicostatus Hall, 1843: 245-246, text-fig. 107(8). 11. Tornoceras (Aulatornoceras) bicostatum (pars). Miller, 1938: 167- Aulatornoceras rhysum (pars). House, 1965: 119-121, pl. 11, fig. 170, pl. 14, fig. 5, pl. 32, fig. 1. 124, text-fig. 13F. Aulatornoceras bicostatum. House, 1962: 262; 1965: 120-121, pl. 10, figs 111-116, pl. 11, figs 125-137, 139-142, text-figs 16A- Type material.–The syntypes were described and illus- D. trated by Clarke and Miller. A lectotype (NYSM 4092) was Truyolsoceras bicostatum. House & Kirchgasser, 1993: 276; Becker, selected by House (1965: 119) who included a specimen 1993a: 218. (BMS E 22464) in the description that is here referred to Aulatornoceras paucistriatum. The type material appears to Type material.–Hall’s type material is AMNH 5888/1, be wholly from the lower Hanover Shale at Java (Wyoming lectotype here designated (Hall, 1843: 245, text-fig. 107 (8); County; Loc. 92). 1879: pl. 72, fig. 9), AMNH 5888/1:1 (Hall, 1879: pl. 72, Discussion.–No new material is assigned here. The type fig. 8, pl. 74, fig. 1), and AMNH 5888/1:2 (Hall, 1879: pl. material leaves much to be desired although there can be lit- 72, fig. 10), all from the Gowanda Shale at Corell’s Point, tle doubt as to the generic assignment. Lake Erie, near Brockton (Chautauqua County). Distribution.–Lower Hanover Shale (West Falls Group; New York material.–As described by House (1965), in- Wyoming County). The type material, all that is recognized, cluding new collections, all from the Gowanda Shale is reported from the lower Hanover Shale, Java (Wyoming (Canadaway Group). County). Dimensions.–See Table 34 and Text-fig. 59B. Range.–Frasnian: Archoceras Genozone UD I-K-?L. Re- Description.–A description based solely on material from gional Zone of Delphiceras cataphractum (23). Conodont the type locality and horizon, using 23 specimens, was pub- Zone MN 12-?13. lished by House (1965), so this species will not be re- described although illustrative material is shown herein. 180 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

Table 34. Biometric data for Truyolsoceras bicostatum (Hall, 1843) from the lower Gowanda Shale. See Text-fig. 59B for graph.

Specimen D WW WH UW WW/WH UW/D

AMNH 5888/1 22.5 — ca. 11.3 —— — (lectotype) AMNH 5888/1:1 ca. 25.0 10.5 12.3 2.8 0.85 0.11 NYSM 6679/B 21.1 8.8 10.5 2.9 0.84 0.14 NYSM 6629C 18.2 8.7 8.5 1.9 1.02 0.10 USNM 137667 ca. 18.0 — ca. 8.8 —— — NYSM 11958 15.3 7.5 8.5 1.9 0.88 0.12 NYSM 6679A 13.7 6.7 7.4 ca. 1.7 0.91 0.12 AMNH 5888/1:2 7.3 — 3.1 1.8 — 0.25 NYSM 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 0.26 NYSM 11962 1.12 0.93 0.62 — 1.50 —

Discussion.–Further statistical information in addition to Known only from the Corell’s Point Goniatite Bed, apart that published in 1965 is shown on the graphs for this from NYSM 12119 and 12120, referred to Truyolsoceras cf. species (Text-fig. 59B); some of this material is from locali- bicostatum, noted above from below that level at Walnut ties for the Corell’s Point Goniatite Bed other than the type Creek (Chautauqua County; Loc. 99a). Noted at most lo- locality at Corell’s Point. Data incorporated in the graphs calities of the Corell’s Point Goniatite Bed between Corell’s (Text-fig. 59B) are based on NYSM 12121-12124, all from Point, Lake Erie Shore (Chautauqua County; Loc. 97), and the Corell’s Point Goniatite Bed in Walnut Creek (Loc. 99b) Clear Creek (North Branch), Marshfield (Erie County; Loc. above the railroad culvert. NYSM 12120 is from 0.9 m (3 106). ft) above the 50-100 mm (2-4 in) siltstone bed that forms Range.–Famennian: Cheiloceras (Cheiloceras) Genozone the base for Mixer Road bridge on the northern side at UD II-C. Regional Zone of Cheiloceras amblylobum (26). Forestville (Chautauqua County; Loc. 99a). NYSM 12119 is from a fallen block at the same locality but thought to be Truyolsoceras clarkei (Miller, 1938) from 3 m (10 ft) above the foundation siltstone. Pl. 30, Figs 1, 3 One of the more noticeable features of this Famennian species is the way in which the ventrolateral furrows are Gephyroceras cf. G. domanicense Clarke, 1904: 345, 380. more decidedly on the outer flanks than in the earlier Frasn- Manticoceras clarkei Miller, 1938: 80-81, pl. 14, figs 15-17. ian forms described here. The species differs substantially Aulatornoceras clarkei. House, 1962: 262; 1965: 121-124, pl. 11, figs 122-123. from Aulatornoceras paucistriatum of the lower Angola Shale Truyolsoceras clarkei. House & Kirchgasser, 1993: 276. (which was mixed with it by Clarke and Miller in their de- scriptions), especially in the much more closed-umbilicate Type material.–The cotypes (NYSM 5652-5654) were early whorls, the asymmetric rather than lingulate lateral described in the works cited above and two of the specimens lobe (Text-figs 60A-C), and the absence of lateral grooves are reillustrated here (Pl. 30, Figs 1, 3). and constrictions and of ventral festoons. The species is Discussion.–This species is like Truyolsoceras bicostatum closer to A. aff. eifliense of this account, but the outer whorls but differs in a wider umbilicus and strongly marked peri- of those specimens are unknown, and outer whorls that are odic ribbing on the flanks. The first impression that these known from the Angola and Hanover Shale specimens seem might be that species in a different preservation is not different. thought to be so, and material in a similar preservation lower All the material is from the Corell’s Point Goniatite Bed in the Gowanda Shale than the Corell’s Point Goniatite Bed apart from two specimens, referred to Truyolsoceras cf. bi- at Forestville is here described as T. cf. bicostatum. costatum, from near the road bridge north of Forestville Distribution:–Gowanda Shale (Canadaway Group; (Loc. 99a) (NYSM 12119 and 12120). These specimens Chautauqua County). Described only from Walnut Creek, have a much more closed umbilicus than the presumably Forestville (Chautauqua County; Loc. 99), but the strata- younger T. clarkei from the Forestville section (compare Pl. graphic level is not known. House (1965) suggested that it 30, Fig. 15, with Pl. 30, Figs 1, 3). might be from the upper Gowanda Shale, Laona Siltstone, Distribution.–Corell’s Point Goniatite Bed, Gowanda or Westfield Shale, but these levels have not been adequately Shale (Canadaway Group; Chautauqua and Erie counties). HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 181

Table 35. Biometric data for Cheiloceras (Cheiloceras) amblylobum (G. & F. Sandberger, 1851) from the lower Gowanda Shale. See Text- fig. 59C for graph.

Specimen D WW WH UW WW/WH UW/D

NYSM 16552 22.7 13.2 13.9 0 0.95 — USNM 137665 14.6 9.1 7.1 ca. 1.0 1.28 0.07 USNM 137666 13.7 ca. 9.0 7.8 ca. 1.0 1.15 0.07 NYSM 16550 13.4 9.3 7.9 ca. 0.5 1.18 0.04 NYSM 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 0.05 NYSM 16553 8.5 6.2 4.8 ca. 0 1.29 — NYSM 16556 6.9 5.9 3.8 0.5 1.55 0.07 NYSM 16555 3.8 2.7 2.4 0 1.12 —

searched in this study. stated that Cheiloceras Trouessart, 1898, had priority, and Range.–Famennian: Cheiloceras (Cheiloceras) Genozone proposed as replacements the names Cheilocerotes and UD II-C. Regional Zone of Truyolsoceras clarkei (25). Cheiloceratos, both of which are invalid (see Richter, 1929: 38); there is also Chiloceras Dreverman (1901: 123), a nom. Superfamily CHEILOCERATOIDEA Frech, 1897 van. As has been elegantly shown by Becker (1993b: 128), (= DIMEROCERATACEAE nom. trans. Bartzsch & this genus plays a major role in the radiation of goniatites Weyer, 1988) following the Upper Kellwasser Event (Text-figs 20-21). The Family CHEILOCERATIDAE Frech, 1897 recognition of the genus in North America was relatively Genus CHEILOCERAS Frech, 1897 late, first in New York, and then in northwestern Canada Type species.–Goniatites subpartitus Münster, 1839, by (House, 1962; House & Pedder, 1963). subsequent designation (Wedekind, 1918: 144). Distribution.–Worldwide except South America and Diagnosis.–Cheiloceratids with conch in earliest stages Antarctica. Marker genus for early Famennian. involute to subevolute, in later stages with closed umbilicus. Range.–Famennian: UD II-B to UD III-C; if Hember- Usually rotund in early stages, later stages can be rotund to gian Raymondiceras are included, UD IV-A; otherwise ex- laterally compressed with venter round to oxyconic; rarely tinct in UD II-G. with gentle ribs, commonly with internal shell varices form- ing constrictions on internal mold. Basic stocks with convex Cheiloceras (Cheiloceras) amblylobum growth lines. Suture EALI to EALU1I. (G. & F. Sandberger, 1851) Discussion.–The question of the name of this group is Pl. 35, Figs 1-10; Text-figs 59C, 61A-B; Table 35 important. No international rules govern taxa above the family level below which adherance to priority is required. Goniatites retrorsus var. amblyloba G. & F. Sandberger, 1851: 108, This is true even of the Code (ICZN, 1999, effective from pl. 10a, figs 20, 24, pl. 10b, fig. 6; 1852: 108. Wedekind, 1918: 146. January 1, 2000). The principle reason for this is to avoid re- Cheiloceras amblylobus. Cheiloceras (Cheiloceras) amblylobum. House, 1962: 274-276; placement of well-known and understood names by the va- Becker, 1993a: 244-246, text-figs 81b, 82e, 83f, pl. 14, figs garies of priority. Frech (1897) established a name group for 10-11; Kirchgasser & House, 1981: 45-46, 49; House & the important genus Cheiloceras that became the best-known Kirchgasser 1993: 276. Famennian goniatite when Wedekind (1918) used it for the Cheiloceras Stufe. The strength of Wedekind’s views are em- Type material.–On the assumption that the originals of phasized by his suborder Cheiloceracea. Proposals to replace the Sandbergers are lost, Matern (1931b) designated a neo- the current group name used here by Dimeroceratacea or type; but House, leaving open the question of loss, chose as Dimeroceratoidea are misconceived and misjudge the im- lectotype the specimen favored by Wedekind (1918: 146) portance of stability in high-taxon terminology, which is the and figured by the Sandbergers (1852: pl. 10, fig. 8). Becker basis of exclusion from priority rulings of all recent Codes. (1993a: 245, text-fig. 83f, pl. 14, figs 10-11) has designated In his detailed review of Nehdenian faunas, Becker a new neotype as Museum für Naturkunde (Berlin), Be (1993a) subdivided this genus into four subgenera: Cheilo- 1256, from the Nehden-Schurbusch, Rhenish Slate Moun- ceras s. s., Raymondiceras, Staffites, and Puncticeras. Only the tains. first is represented in New York. Strand (1929: 8) wrongly New York material.–Fifteen pyritic molds were listed pre- 182 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

imens in the collections of MRH now deposited with the NYSM 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 need not be repeated. Statistical details of additional material is included in Table 35 and Text-fig. 59C. Typical sutures are illustrated in Text-figs 61A-B, and a range of specimens are illustrated 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), Walnut Creek (Loc. 99), and Smith Mills (Loc. 100). From Catta- raugus County: Big Indian Creek (Loc. 101), Little Indian Creek (Loc. 102), Cattaraugus Creek (Cattaraugus Indian Reservation) (Loc. 103), and Cattaraugus Creek (South Branch) (Loc. 104). From Erie County: Clear Creek (North Branch), 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 locate the level. Range.–Famennian: Cheiloceras (Cheiloceras) Genozone UD 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 should Text-fig. 61. Sutures of Cheloceras, Maeneceras, and Sporadoceras be referred to as Sporadoceratoidea (nom. trans. Miller & spp. from New York, Pennsylvania, and Ohio (after House, 1962, Furnish, 1957) rather than Praeglyphioceratoidea and House et al., 1986). A-B. C. (Cheiloceras) amblylobum (G. & F. Sandberger, 1851). A. USNM 137665, suture based on speci- (Ruzhencev, 1957), as in Becker (1993a), to conform with men collected by W. Moran from the Gowanda Shale at Corell's the ICZN guidance concerning the least disruption of senior Point (Loc. 97), on the shore of Lake Erie, Chautauqua County, taxon names. The group comprises forms with convex suture reversed, X 3. B. USNM 137666, suture based on specimen growth lines and sutures that differ from the Cheiloceratidae from same locality as A, X 2.8. C-D. aff. (G. & F. M. acutolaterale in the addition of extra adventitious ventral lobes (A2 and in Sandberger, 1850), USNM 137645, based on a specimen collected some A3). by H. S. Williams in 1884 from the Ellicott Shale at Porter's Creek, Summerdale, Chautauqua County. C. Suture at 83 mm di- Genus MAENECERAS Hyatt, 1884 ameter, X 0.7. D. Reversed suture at 117 mm diameter, a com- Type species.–Goniatites acutolateralis G. & F.Sandberger, posite diagram, X 0.7. E. M. milleri (Flower & Caster, 1935), 1850, by original designation (Hyatt, 1884: 321). diagram of the suture (reversed), from Miller (1938) of the holo- type from Howard Quarry, Erie County, Pennsylvania, X 0.7. F. Diagnosis.–Sporadoceratids with discoidal to subglobular M. inflexum (Wedekind, 1908), NMNH 240513, suture based on conchs with closed umbilicus and convex growth lines. Su- pyritic fragments from 2 m above the base of the Cleveland Shale, ture EA2A1LUI but with A1 sharp or narrowly rounded and Birmingham, Ohio, X 2.5. A2 shallow and rounded. For discussion and included species, see Becker (1993a). viously (House, 1962: 276), including USNM 137665 and Distribution.–Widespread. North America (very rare), 137666 collected by W. Moran, NYSM 11239-11242 col- Europe, Russia, Asia, North Africa, Australia, and South lected by J. M. Clarke and D. D. Luther in 1898, and spec- America. HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 183

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 small and the extent to which the A2 lobe would deepen in later stages is uncertain. The specimen can be compared to the holotype of Maeneceras acutolaterale, from which it differs in the more parallel-sided conch and in the lesser depth of the A2 lobe, but the A1 lobe is not so sharp. Distribution.–Ellicott (Chadakoin) Shale (Conneaut Group; Chautauqua County). The specimen is labeled HSW coll. 518B. J. W. Wells informed MRH that locality 518B is marked on manuscript maps of H. S. Williams, then at Cornell University, which showed it to be along Porter 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. Text-fig. 62. Maeneceras aff. acutolaterale (G. & F. Sandberger, 110 of this report), This point lies in the middle part of the 1850), USNM 137645, collected by H. S. Williams in 1884 from outcrop of the Ellicott (Chadakoin) Shale on the map of the Ellicott Shale at Porter's Creek, Summerdale, Chautauqua Tesmer (1954: 29). County. Scale = 10 cm. Range.–In New York: Famennian: Maeneceras Genozone UD II-G. Regional Zone of M. aff. acutolaterale (27). In Range.–Famennian: UD II-G to V-B (Becker, 1993b). Germany, the type species is said to range from ?upper biferum Zone (UD II-G) to the contiguum Zone (UD II-H) Maeneceras aff. acutolaterale (G. & F. Sandberger, 1850) (Becker, 1993a: 314). In Pennsylvania, the occurrence of M. Pl. 35, Figs 11, 13; Text-figs 61C-D, 62 milleri (illustrated here, Text-fig. 61E) higher in the North American succession, in the Conewango Group (Cattarau- aff. Goniatites acutolaterale G. & F. Sandberger, 1850: pl. 6, figs 1, gus-Oswayo equivalents), Howard Quarries ( Loc. 111), Re- 1a-b; 1851: 98. gional Fauna 28, correlates with UD II-H (Becker & House, aff. Maeneceras acuto-laterale. Hyatt, 1884: 321. 2000: 134). Still higher, in Ohio, M. inflexum (Text-fig. Sporadoceras cf. pompeckji Wedekind, 1918. House 1962: 276, pl. 61F) from the Cleveland Shale (Regional Fauna 29), corre- 46, fig. 12, text-fig. 12A. lates with Prionoceras Genozone UD IV-A (Becker & House, Maeneceras cf. pompeckji. Kirchgasser & House, 1981: 46, 49; 2000). House & Kirchgasser, 1993: 276. Family and Genus Indeterminable Type material.–Holotype of Maeneceras acutolaterale de- scribed by the Sandberger brothers, refigured by Becker ?Genus edwinhalli (Clarke, 1898) (1993a: 313, fig. 96). Pl. 35, Fig. 12 New York material.–One specimen only (USNM 137645) collected by H. S. Williams in 1884 from Porter Tornoceras Edwin-halli Clarke, 1898, 1899a, b: 111, text-fig. 85. Creek, Summerdale (Chautauqua County). Tornoceras (Tornoceras) edwinhalli. Miller, 1938: 151, pl. 31, figs Dimensions.–USNM 137645, maximum D = 132 mm, 10-11. D = 120 mm, WW = 69 mm, and UW = ?0 mm. Description.–Features of shell as in Text-fig. 62. Body Discussion:–This single specimen (NYSM 4090) de- chamber occupying last one-third whorl; phragmocone scribed by both Clarke and Miller needs no further descrip- showing sutural approximation. Laterally compressed and tion. However, it seems that the drawn figure is not a ventral discoidal with rounded venter and closed umbilicus. A2 lobe suture at all, but the impression of the dorsal suture from a showing increasing depth in last half whorl of phragmocone. whorl now lost. Such simple dorsal sutures are found in Growth lines not seen. some prolobitids, but in the absence of further information, Discussion.–At present, the distinction between and perhaps the dissection of the specimen, no more precise Maeneceras and Sporadoceras lies in the depth of the A2 lobe. naming is possible. Yet diagnoses do not state the percentage of depth required, Distribution.–Conneaut Group, Nile (Allegany County). nor the shell diameter at which this must be attained. The Range.–Famennian. 184 BULLETINS OF AMERICAN PALEONTOLOGY no. 374 HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 185

Text-fig. 63 (at left and above). Diagramatic illustrations of zone-defining goniatites in the late Devonian of New 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. Chutoceras nundaium (Text-figs 28I-J); 16a. Koenenites styliophilus styliophilus (Text-figs 34F,L); 16b. K. styliophilus kilfoylei n. 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. Prochorites alveolatus (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. Crickites lindneri (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. 186 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

SYNOPSIS Text-fig. 63 summarizes the succession of zone-defining go- ACKNOWLEDGMENTS niatites in the late Devonian of New York State with illus- Progress on this work has proceeded over many years and trations of sutures and whorl-sections of some key taxa. consequently we are indebted to many for their help and as- sistance. The Natural Environment Research Council ADDENDUM (NERC) in the 1960s funded a research project to MRH Noted here are recent reports concerning some of the taxa that enabled us both to commence a systematic study of the and sections described in this monograph. Work et al. Late Devonian across the State of New York during which (2007) reported the discovery of Pharciceras in the New Al- time stratigraphical successions were established in a general bany Shale (Trousdale Member) in Kentucky. Pharciceras way and systematic collections begun. The New York State barnetti Work et al., 2007, is similar to P. amplexum from Museum and Science Service funded field work at that time the Upper Tully Limestone but differs in having a wider by WTK and provided a vehicle for two seasons’ work and, conch and more depressed whorl form. The two horizons through the kind offices of its staff, helped in many ways. with P. barnetti correlate to an interval within the lower We thank especially L. V. Rickard for putting his enormous Geneseo Shale of the Taghanic Onlap sequence that spans knowledge of the Devonian geology of the State at our dis- the level of the Fir Tree Limestone (and equivalent Leicester posal; D. W. Fisher, former State Paleontologist, who facil- Pyrite) of the upper Givetian (MD III A; hermanni and dis- itated work in every way; and C. Kilfoyle for guidance and paralis conodont zones). loans from the outstanding State collections at Albany, and Baird et al. (2006a) have greatly refined the sections of for permission to photograph and access the State collec- the lower to middle Genesee Group between Lake Erie and tions. More recently, E. Landing and L. Van Aller Hernack the Genesee Valley. The delineation of the many discontinu- have provided access to the collections and C. A. Ver ities, and particularly those associated with the Lodi and Straeten has given bibliographic help. At the University of Linden Horizons and the Lower and Upper Divisions of the Rochester, R. G. Sutton graciously provided data on black Genundewa Limestone, should lead to an even finer-scale shale correlations that proved to be crucial in determining resolution of the goniatite succession. the Rhinestreet equivalent goniatite sequences, and G. Kloc Zambito et al. (2007, unpublished data) have revised has generously shown us his collections. Jean Dougherty of and analyzed the Sherburne, Renwick, and Ithaca Forma- the Geological Survey of Canada has advised on the material tions and equivalents of the Genundewa Limestone and collected by E. M. Kindle. West River Shale in the Ithaca area using the methodology To Cornell University, and the late Professor J. W. Wells of sequence stratigraphy. Newly measured sections include in particular, we owe a debt that cannot be repaid in words. the classic section at Fall Creek, Ithaca. The goniatite succes- John Wells introduced both of us to the geology of central sion outlined herein is illustrated in a composite section of New York and supervised the doctorate by WTK on the the lower, middle, and part of the upper Genesee Group. Cashaqua Shale. Throughout the period until his untimely Several submember and system-tract units are tentatively death, he gave his whole-hearted encouragement and put correlated with the Genesee Group succession to the west. his encylopedic knowledge and substantial collections of fos- The new sequence stratigraphy provides a framework for re- sils, books, and maps at our disposal. With his marvellous fining the ranges of genera and species of goniatites in the wife, Pie Wells, he provided respite and refreshment count- Ithaca region. less times during fieldwork and odd visits to their home, Finally, Baird et al. (2006b) reported the discovery of Lucky Stone Lodge, at Sheldrake on the banks of Cayuga Naplesites in the lower Rhinestreet Shale at Lake Erie shore. Lake. The occurrence is a bedding plane surface in black shale The Paleontological Research Institution (PRI) at Ithaca with numerous flattened specimens (some partially pyritized has provided material for study, as has the Buffalo Museum. and showing sutures) and a few uncrushed specimens in There was a major early input by the U. S. National Mu- scattered concretions at the same level. The species is likely seum when G. A. Cooper introduced MRH to many aspects to be either N. inyx or N. naplesense, the species described by of North American Devonian work, and material in the Mu- Clarke (1898) from an unknown level around Naples, On- seum collections has been made available for study. The help tario County. The discovery of Naplesites at Lake Erie con- of others at the Museum, especially W. A. Oliver, Jr., Helen firms our guess that Clarke’s specimens came from the lower Duncan, Jean Berdan, and J. T. Dutro, Jr., is much appre- Rhinestreet Shale. Work has begun to see if the Naplesites ciated. At the American Museum of Natural History, New horizon at Lake Erie can be located in the section at Naples. York, Otto Haas and N. H. Landman have facilitated access -WTK to collections, as have N. Nitecki and S. Lidgard at the Field HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 187

Museum of Natural History in Chicago. At the PRI, W. D. August 2002) in Dorchester near his home in Weymouth, Allmon and W.Taylor have facilitated our work. At the Uni- Dorset, England. The task of editing this final manuscript versity of Iowa, B. F. Glenister and W. M. Furnish have thus fell to me (WTK). Some parts needed updating but I made Devonian comparative material available and, over the have made every effort to keep the content and style as close last decade, collaboration with the conodont biostrati- as possible to the original. Thanks to my new colleagues at graphic studies of G. Klapper has sharpened aspects of our the Department of Geology for supporting the project in work. More recently we have appreciated demonstration of various ways: M. Rygel, C. Kelson, and R. Greene (Secre- their work on Devonian rocks in New York by C. E. Brett, tary). Thanks also to Betsy Northrop for twenty-fve years of G. Baird, and D. J. Over. We are both greatly indebted for support and patience. Finally, special thanks to Nancy Dutro astute comments and help by our colleague, R. T. Becker, of for a fine job of copy-editing for PRI and to Paula the University of Münster, Germany, for his collaboration in Mikkelsen, Director of Publications at PRI, for the final ed- fieldwork in New York as well as in Australia, Russia, Eu- iting and putting the parts together for publication. All er- rope, and North Africa, and for his systematic collation of rors remaining are of course my responsibility. literature that he has put freely at our disposal. Michael House and I agreed long ago to dedicate this This work has continued while MRH was successively a monograph to the memory of Professor and Mrs. John West member of Geology Departments at Oxford, Hull, and Wells. Southampton Universities, where staff provided secretarial and photographic help; J. Garner and B. A. Marsh provided LITERATURE CITED most of the prints used on the plates. Many thanks are due to Felicity House for continuing support. Aboussalam, Z. S. 2003. Das “Taghanic-Event” im höheren At the State University of New York at Potsdam, WTK Mitteldevon von West-Europa und Marokko. Mün- has been supported since 1969 by numerous grants from sterche Forschungen zur Geologie und Paläontologie, 97: 1- the SUNY Research Foundation, the New York State United 330. University Professions, and Potsdam College. The related Aboussalam, Z. S., & R. T. Becker. 2001. Prospects for an collaboration on conodonts with G. Klapper at the Univer- upper Givetian substage. Mitteilungen aus dem Museum sity of Iowa was supported by National Science Foundation für Naturkunde im Berlin,Geowissenschaftiche, Reihe, 4: Research Opportunity Award (EAR-890475) to WTK. 83-99. Thanks also are due to the many undergraduate students Ager, D, V. 1967. Brachiopod palaeoecology. Earth Science who assisted WTK on the conodont work; they are ac- Review, 3: 157-179. knowledged by name in separate publications. The generous Anonymous. 1993. New York State Atlas and Gazeteer, 4th support over the years of many colleagues at Potsdam Col- ed. DeLorme Mapping, Freeport, Maine, 104 pp. lege is greatly appreciated, particularly the staffs of the Office Archiac, E. J. A. D. d’, & M. E. de Verneuil. 1842. On the of Scholarships and Grants, Crumb Library, and Computer fossils of the older deposits in the Rhenish provinces, Services; special thanks go also to R. Bitely for his advice preceded by a general survey of the fauna of the Paleo- and help on photographic work. We are also indebted to our zoic rocks and followed by a tabular list of the organic re- friends on the faculty and staff of the SUNY Potsdam Ge- mains of the Devonian System in Europe. Transactions of ology Department, Professors F. Revetta, B. Van Diver, N. the Geological Society of London, series 2, 6: 303-410, pls O’Brien, J. Carl, and R. Badger, J. Chiarenzelli, L. Amati 25-38. and the Departmental Secretaries, J. Moriarty, V. O’Brien, Arthaber, G. von. 1911. Die Trias von Albanien. Beiträge and J. Bullis, for their years of support and encouragement. zur Paläontologie und Geologie Österreich-Ungarns und Thanks especially to J. Bullis, for preparing the final type- des Orients, 24: 169-277. script, and Patty Stone for much technical assistance. Baird, G. C. 1976. Coral encrusted concretions: a key to Thanks also to Kim Coleman for computer-drafting Text- recognition of a 'shale-on-shale' erosion surface. Lethaea, figs 1 and 63. 9: 293-302. We greatly appreciate the timely reviews of the manu- Baird, G. C. 1979. Sedimentary relationships of Portland script provided by Gordon Baird (SUNY Fredonia), Brian Point and associated Middle Devonian rocks in central Glenister (University of Iowa), and Warren Allmon (Pale- and western New York. New York State Museum Bulletin, ontological Research Institution); most of their suggestions 433: 1-24. have been incorporated in this final version. A few days after Baird, G. C., & C. E. Brett. 1986a. Erosion of an aerobic receiving the reviews and notification of acceptance for pub- seafloor: significance of reworked pyrite deposits from lication of the manuscript, Michael House passed away (06 the Devonian of New York state. Palaeogeography, Palaeo- 188 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

climatology, Palaeoecology, 57: 157-193. state. Pp A1-A45, in: Western New York and Ontario, Baird, G. C., & C. E. Brett. 1986b. Submarine erosion on Field Trip Guidebook, New York State Geological Associa- the dysaerobic sea floor: Middle Devonian corrasional tion, 62nd Annual Meeting, G. G. Lash (ed.), Depart- disconformities in the Cayuga Valley region. Pp 23-80, ment of Geosciences, State University of New York, in: Field Trip Guidebook, New York State Geological Asso- Fredonia. ciation, 58th Annual Meeting, Cornell University, Ithaca, Baird, G. C., & R. Jacobi. 1999. “Nunda Sandstone” depo- New York, Hempstead, New York. sitional event in the Pipe Creek black shale, South Wales- Baird, G. C., & C. E. Brett. 2001. Eustatic and fluxural Varysburg area, New York. Pp B1-B7, in: Field Trip events recorded in the late Middle Devonian Tully For- Guidebook, New York State Geological Association, 71st mation, New York state and Pennsylvania. Geological So- Annual Meeting, G. G. Lash (ed.), Department of Geo- ciety of America, Abstracts with Programs, 33(1): A17. sciences, State University of New York, Fredonia. Baird, G. C., & C. E. Brett. 2003. Shelf and off-shore de- Baker, C., B. F. Glenister, & C. O. Levorson. 1986. Devon- posits of the Tully Formation in New York and Pennsyl- ian ammonoid Manticoceras from Iowa. Proceedings of vania: faunal incursions, eustacy and tectonics. Pp the Iowa Academy of Science, 93 (1): 7-15. 141-156, in: Mid-Paleozoic Bio- and Geodynamics: the Bartzsch, K., & D. Weyer. 1988. Die unterkarbonische Am- North Gondwana-Laurussia Interaction, P. Königshof & monoidea-Subfamilia Karagandoceratinae. Freiberger E. Schindler (eds), Proceedings of the 15th International Forschungsheft, C419: 130-412. Senckenberg Conference, Courier Forschungsinstitut Becker, R. T. 1986. Ammonoid evolution before, during Senckenberg, 242. and after the “Kellwasser-event” – review and prelimi- Baird, G. C., C. E. Brett, & A. J. Bartholomew. 2003. Late nary new results. Pp 181-188, in: Global Bio-Events; a Middle Devonian biotic and sedimentary events in east Critical Approach, O. H. Walliser (ed.), Springer-Verlag, central New York-Tully Formation clastic correlative suc- Berlin and New York. cession in the Sherburne-Oneonta area. Pp 1-54, in: Becker, R. T. 1990. Stratigraphische Gliederung und Am- Field Trip Guidebook, New York State Geological Associa- monoideen Fauna im Nehdenium (Oberdevon II) von Eu- tion, 75th Annual Meeting, E. L. Johnson (ed.), Geology rope und Nord Africa. Unpublished Ph. D. Dissertation, and Environmental Science Department, Hartwick Col- University of Bochum, Bochum, Germany, 464 pp, 26 lege, and Department of Earth Sciences, State University pls. of New York, College at Oneonta, Oneonta, New York. Becker, R. T. 1993a. Stratigraphische Gliederung und Am- Baird, G. C., C. E. Brett, & W. T. Kirchgasser. 1989. Gen- monoideen-Faunen im Nehdenium (Oberdevon II) von esis of black shale-roofed discontinuities in the Devonian Europa und Nord Afrika. Courier Forschungsinstitut Genesee Formation, western New York state. Pp 357- Senckenberg, 155: 1-405, 26 pls. 375, in: Devonian of the World, Proceedings of the Second Becker, R. T. 1993b. Anoxia, eustatic changes, and Upper International Symposium on the Devonian System, Calgary, Devonian to lowermost Carboniferous global am- vol. 2, Sedimentation, N. J. McMillan, A. F. Embry, & J. monoid diversity. Pp 115-163, in: The Ammonoidea: En- D. Glass (eds), Canadian Society of Petroleum Geolo- vironment, Ecology and Evolutionary Change,M.R. gists Memoir 14. House (ed.), Systematics Association Special Volume 47, Baird, G. C., W. T. Kirchgasser, D. J. Over, & C. E. Brett. Clarendon Press, Oxford, UK. 2006a. Middle-Upper Devonian depositional and biotic Becker, R. T. 1995. Taxonomy and evolution of Late Fa- events in western New York. Pp 128-164, in: New York mennian Tornocerataceae (Ammonoidea). Berliner Ge- State Geological Association, 78th Annual Meeting, Octo- owissenschaftliche, Abhandlungen, E-16: 607-643. ber 6-8, 2006, R. D. Jacobi (ed.). New York State Geo- Becker, R. T., & M. R. House. 1993. New early Upper De- logical Association, Buffalo, New York. vonian (Frasnian) goniatite genera and the evolution of Baird, G. C., W. T. Kirchgasser, D. J. Over, & C. E. Brett. the "Gephurocerataceae." Berliner Geowissenschaftliche, 2006b. An early late Devonian bone-bed-pelagic lime- Abhandlungen, E- 9: 111-133, 2 pls. stone succession: the North Evans-Genundewa Lime- Becker, R. T., & M. R. House. 1994a. Kellwasser events and stone story. Pp 354-395, in: New York State Geological the goniatite succession in the Devonian of the Mon- Association, 78th Annual Meeting, October 6-8, 2006, R. tagne Noire with comments on possible causations. D. Jacobi (ed.). New York State Geological Association, Courier Forschungsinstitut Senckenberg, 169: 45-77. Buffalo, New York. Becker, R. T., & M. R. House. 1994b. International De- Baird, G. C., & G. G. Lash. 1990. Devonian strata and pa- vonian goniatite zonation, Emsian to Givetian, with new leoenvironments: Chautauga County region: New York records from Morocco. Courier Forschungsinstitut Senck- HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 189

enberg, 169: 79-135. Present, Vienna, Austria, September 6-9, 1999. Geolo- Becker, R. T., & M. R. House. 1997. Sea-level changes in gische Bundesanstalt, 57: 279-297. the Upper Devonian of the Canning Basin, Western Boekenkamp, R. P. 1963. The Stratigraphy of the Sherburne Australia. Pp 129-146, in: On Sea-level Fluctuations in Member (Genesee Formation ) of the Cayuga Trough, Cen- the Devonian, M. R. House & W. Ziegler (eds). Courier tral New York. Unpublished M. S. Thesis, Cornell Uni- Forschungsinstitut Senckenberg, 199. versity, Ithaca, New York, 42 pp. Becker, R. T., & M. R. House. 2000. Devonian ammonoid Bogoslovsky, B. I. 1955a. On the family Pharciceratidae zones and their correlation with established series and Hyatt, 1900 (in Russian). Doklady Akademiya Nauk stage boundaries. Pp 113-151, in: Subcommission on De- SSSR, 103: 1103-1106. vonian Stratigraphy, Fossil Groups Important for Boundary Bogoslovsky, B. I. 1955b. Devonian ammonoids from the Definition, P. Bultynek (ed.). Courier Forschungsinstitut Rudny Altai (in Russian). Bulletin Imperialskogo Senckenberg, 220. Moskovskago Obshchestva Ispytatelei Prirody, Otdel Ge- Becker, R. T., & M. R. House. 2000a. Late Givetian and ologiya, 30: 94-95. Frasnian ammonoid succession at Bou Tchrafine (Anti Bogoslovsky, B. I. 1957. New genera of Devonian am- Atlas, southern Morocco). Notes et Mémoires de Service monoids (in Russian). Materialy k Osnovam Paleon- Géologique de Maroc (Rabat), 399: 27-36. tologii, Akademiya Nauk SSSR, Paleontologicheskii Becker, R. T., & M. R. House. 2000b. The Famennian am- Institut, 1: 45-48. monoid succession at Bou Tchrafine (Anti Atlas, south- Bogoslovsky, B. I. 1958. Devonian Ammonoidea of the ern Morocco). Notes et Mémoires de Service Géologique Rudny Altai (in Russian). Trudy Paleontologicheskogo In- de Maroc (Rabat), 399: 37-42. stituta, Akademiya Nauk SSSR, 64: 1-155, pls 1-9. Becker, R. T., M. R. House, & W.T. Kirchgasser. 1993. De- Bogoslovsky, B. I. 1965. Drevneyshiy, zhivetskiy predstavitel vonian biostratigraphy and timing of facies movements podotryada Gephuroceratina. Paleontologicheskiy Zhur- in the Frasnian of the Canning Basin, Western Australia. nal, 1965(2): 50-54. Pp 293-321, in: High Resolution Stratigraphy, E. A. Hail- Bogoslovsky, B. I. 1969. Devonian Ammonoids, I. Agoni- wood & R. B. Kidd (eds), Geological Society Special atitida (in Russian). Trudy Paleontologicheskogo Instituta, Volume 70, London. Akademiya Nauk SSSR, 124: 1-341, pls 1-29. Becker, R. T., M. R. House, W.T. Kirchgasser, & P.E. Play- Bogoslovsky, B. I. 1971. Devonian Ammonoids, II. Goni- ford. 1991. Sedimentary and faunal changes across the atitida. Trudy Paleontologicheskogo Instituta, Akademiya Frasnian/Famennian boundary in the Canning Basin of Nauk SSSR, 127: 1-228, pls 1-19. Western Australia. Historical Biology, 5: 183-196. Bowen, Z. P.,D. C. Rhoads, & A. L. McAlester. 1974. Ma- Becker, R. T., M. R. House, V. V. Menner, & N. S. Ov- rine benthic communities in the Upper Devonian of natanova. 2000. Revision of ammonoid biostratigraphy New York. Lethaia, 7: 93-120. in the Frasnian (Upper Devonian) of the Southern Bradley, W. H., & J. F. Pepper. 1938. Structure and gas pos- Timan (Northeast Russian Platform). Geologica Polonica, sibilities of the Oriskany sandstone in Steuben, Yates, 50(1): 67-97. and parts of the adjacent counties, Pt. 1 of Geologic Bensaïd, M. 1974. Étude sur des goniatites à la limite du structure and occurence of gas in parts of southwestern Devonien moyen et supérieur, du Sud Morocain. Notes New York. United States Geological Survey Bulletin, 899A: et Mémoires de Service Géologique du Maroc, 36(264): 81- 1-68. 140, pls 1-6. Brett, C. E., & G. C. Baird. 1982. Upper Moscow-Genesee Bergin, M. J. 1964. Bedrock geology of the Penn Yan and stratigraphic relationships in western New York: evi- Keuka Lake quadrangles, New York. United States Geo- dence for regional erosive beveling in the late Middle logical Survey Bulletin, 1161 G: 1-34. Devonian. Pp 19-63, in: Geology of the Northern Ap- Beyrich, E. 1837. Beiträge zur Kenntniss der Versteinerungen palachian Basin, Guidebook for Field Trips in Western New des Rheinischen Übergansgebirges, 1. Berlin, iv + 44 pp. York, Northern Pennsylvania and Adjacent Southern On- Beyrich, E. 1884. Erlauterungen zu den Goniatiten L. V. tario, New York Geological Association, 54th Annual Meet- Buch's. Zeitschrift der Duetschen Geologische Gesellechaft, ing, E. J. Buehler & P. E. Calkin (eds), State University 36: 203-219. of New York at Buffalo, Buffalo, New York. Bockwinkel, J., R. T. Becker, & V. Ebbighausen. 1999. Mor- Brett, C. E., & G. C. Baird. 1985. Carbonate-shale cycles in phometry and taxonomy of lower Famennian Sporado- the Middle Devonian of New York: an evaluation of ceratidae (Goniatitida) from southern Morocco. Fifth models for the origin of limestones in terrigenous shelf International Symposium on Cephalopods, Past and sequences. Geology, 13: 324-327. 190 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

Brett, C. E., & G. C. Baird. 1986. Submarine erosion on the Guide, International Union of Geological Sciences, Sub- dysaerobic seafloor: Middle Devonian corrasional dis- commission on Devonian Stratigraphy, July 22-27, 1997. conformities in the Cayuga Valley region. Pp 23-80, in: Department of Earth and Environmental Sciences, Uni- Fieldtrip Guidebook, New York Geological Association, versity of Rochester, Rochester, New York, 369 pp. 58th Annual Meeting, Cornell University, Ithaca, New Brown, T. 1841. Description of some new species of fossil York, Hempstead, New York. shells, found chiefly in the vale of Todmorden, Yorkshire. Brett, C. E., & G. C. Baird. 1990. Submarine erosion and Transactions of the Manchester Geological Society, 1: 212- condensation in a foreland basin: examples from the De- 229. vonian of Erie County, New York. Pp A1-A56, in: West- Buch, L. von. 1832. Über Goniatiten. Physikalische ern New York and Ontario, Field Trip Guidebook, New Akademie Wissenschaft Berlin, Abhandlungen, 1930: 159- York State Geological Association, 62nd Annual Meeting, 187, pls 1-5. G. G. Lash (ed.), State University of New York, Fredo- Buggisch, W. 1972. Zur Geologie und Geochemie der Kell- nia. wasserkalke und der Begleitsedimente (Unteres Oberde- Brett, C. E., & G. C. Baird. 1994. Depositional sequences, von). Hessisches Landesamt für Bodenforschung. cycles, and foreland basin dynamics in the late Middle Abhandlungen, 62: 1-68. Devonian (Givetian) of the Genesee Valley and western Buggisch, W., & C. D. Clausen. 1972. Conodonten- und Finger Lakes region. Pp 505-568, in: Field Trip Guide- Goniatiten-Faunen aus dem oberen Frasnium und un- book, New York State Geological Association, 66th Annual teren Famennium Marokkos (Tafilalt, Antiatlas). Neues Meeting, C. E. Brett & J. Scatterday (eds), University of Jahrbuch für Geologie und Paläontologie, Abhandlungen, Rochester, Rochester, New York. 141: 137-167. Brett, C. E., & G. C. Baird. 1996. Middle Devonian sedi- Caster, K. E. 1933. Upper Devonian rocks at Ithaca, New mentary cycles and sequences in the northern Ap- York (abstract). Geological Society of America Bulletin, 44: palachian Basin. Pp 213-239, in: Paleozoic Sequence 201-202. Stratigraphy: Views from the North American Craton, B. J. Chadwick, G. H. 1920. Large fault in western New York. Witzke, G. A. Ludvigson, & J. Day (eds), Geological So- Geological Society of America Bulletin, 31: 117-120. ciety of America Special Paper 306, Boulder, Colorado. Chadwick, G. H. 1933. Great Catskill delta: and revision Brett, C. E., & G. C. Baird. 1997. Middle Devonian stratig- of Late Devonic succession. Pan American Geologist, 60: raphy, facies, and depositional environments of western 91-107. New York State. Pp 67-78, in: Devonian Cyclicity and Chadwick, G. H. 1935a. Faunal differentiation in the Sequence Stratigraphy in New York State, Field Trip Guide- Upper Devonian. Geological Society of America Bulletin, book, International Union of Geological Sciences, Subcom- 46: 305-342. mission on Devonian Stratigraphy, July 22-27, 1997, C. Chadwick, G. H. 1935b. Chemung is Portage. Geological E. Brett & C. A. Ver Straeten (eds), Department of Society of America Bulletin, 46: 343-354. Earth and Environmental Sciences, University of Chao, K. 1956. Notes on some Devonian ammonoids from Rochester, Rochester, New York. Southern Kwangsi. Acta Palaeontologica Sinica, 4: 101- Brett, C., G. C. Baird, & W. T. Kirchgasser. 1997. Stop A- 116. 5. Lake Erie cliffs south of Eighteenmile Creek. Pp 172- Clarke, J. M. 1882. New phyllopod crustaceans from the 179, in: Devonian Cyclicity and Sequence Stratigraphy in Devonian of western New York. American Journal of Sci- New York State, Field Trip Guidebook, International ence, series 3, 13: 476-478, pl. 1. Union of Geological Sciences, Subcommission on Devonian Clarke, J. M. 1883. New discoveries in Devonian Crustacea. Stratigraphy, July 22-27, 1997, C. E. Brett & C. A. Ver American Journal of Science, 25: 120-125. Straeten (eds), Department of Earth and Environmental Clarke, J. M. 1884. Über deutsch oberdevonische Crus- Sciences, University of Rochester, Rochester, New York. taceen. Neues Jahrbuch für Minerogie, Geologie und Brett, C. E., G. C. Baird, & C. A. Ver Straeten. 1999. Flex- Paläontologie, 1: 178-185, pl. 5. uval events and faunal changes preceding the third Aca- Clarke, J. M. 1885. On the higher Devonian faunas of On- dian tectophase in the late Givetian of the northern tario County, New York. United States Geological Survey Appalachian Basin region. Pp 4-5, in: Abstract Book, Er- Bulletin, 16: 1-60. rachidia Meeting, SDS-IGCP 421, Faculty of Sciences, Clarke, J. M. 1891a. The fauna with Goniatites intumescens, Moulay Ismail University, Errachidia, Morocco. Beyrich, in western New York. American Geologist, 8: 86- Brett, C., & C. A. Ver Straeten, eds. 1997. Devonian Cyclic- 105. ity and Sequence Stratigraphy in New York State, Field Trip Clarke, J. M. 1891b. Die Fauna mit Goniatites intumescens HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 191

im westlichen New York. Neues Jahrbuch für Mineralogie, Sicht. Neues Jahrbuch für Geologie und Paläontologie, Ab- Geologie und Paläontologie, 1: 161-168. handlungen, 137: 175-208. Clarke, J. M. 1892. The discovery of Clymenia in the fauna Coates, D. R. (ed.). 1963. Geology of South-Central New of the Intumescens Zone (Naples Beds) of western New York. Guidebook, New York State Geological Associa- York and its geological significance. American Journal of tion, 35th Annual Meeting, Binghamton, New York, Science, series 3, 43: 57-63. 116 pp. Clarke, J. M. 1897. The stratigraphic and faunal relations of Cole, W. S., et al. 1959. Geology of the Cayuga Basin, 2nd the Oneonta sandstones and shales, the Ithaca and (rev.) ed. Guidebook, New York State Geological Associ- Portage groups in central New York. New York State Ge- ation, 31st Annual Meeting, Cornell University, Ithaca, ologist Annual Report, 15: 11-12, 27-81. New York, 36 pp. Clarke, J. M. 1898. The Naples Fauna (Fauna with Mantic- Colton, G. W. 1956. Bedrock geology of the Hamburg oceras intumescens) in Western New York (preprint valid Quadrangle, New York. United States Geological Survey for taxonomic purposes). Albany, New York, i + 31-161 Map 9097. pp, pls 1-9. Colton, G. W., & W. de Witt, Jr. 1958. Stratigraphy of the Clarke, J. M. 1899a. The Naples fauna (fauna with Mantic- Sonyea Formation of Late Devonian age in western and oceras intumescens) in western New York (transmitted to west-central New York. United States Geological Survey, the Legislature April 20, 1897). Sixteenth Annual Report Oil and Gas Investigation Chart OC-54. of the [New York] State Geologist for the Year 1896: 29- Conrad, T. A. 1842. Observations on the Silurian and De- 161, pls 1-9. vonian systems of the United States, with descriptions Clarke, J. M. 1899b. The Naples fauna (fauna with Mantic- of new organic remains. Journal of the Academy of Natu- oceras intumescens) in western New York. New York State ral Sciences of Philadelphia, 8: 228-280. Museum, 50th Annual Report of the Regents, 1896. 2, Re- Conybeare, W. D., & W. Phillips. 1822. Outlines of the Ge- port of the State Geologist and his Field Assistants. Univer- ology of England and Wales. W. Phillips, London, 470 pp. sity of the State of New York: 29-161, pls 1-9. Cooper, G. A. 1930. Stratigraphy of the Hamilton group of Clarke, J. M. 1899c. Guide to the fossiliferous rocks of New New York. American Journal of Science, series 5, 19: 116- York state. University of the State of New York, Handbook, 134, 214-236. 15: 1-110. Cooper, G. A. 1933. Stratigraphy of the Hamilton Group of Clarke, J. M. 1902. The indigene and alien faunas of the eastern New York. American Journal of Science, series 5, New York Devonic. New York State Museum Bulletin, 52: 26: 537-551. 664-672. Cooper, G. A. 1968. Age and correlations of the Tully and Clarke, J. M. 1903. Classification of the New York series of Cedar Valley Formations in the United States. Pp 701- geologic formations. New York State Museum Handbook, 709, in: International Symposium on the Devonian System, 19: 1-28. Calgary, 1967, vol. 2, D. H. Oswald (ed.), Alberta Soci- Clarke, J. M. 1904. Naples fauna (fauna with Manticoceras ety of Petroleum Geologists, Calgary, Alberta, Canada. intumescens), Part 2. New York State Museum Memoir, 6: Cooper, G. A., & J. S. Williams. 1935. Tully Formation of 31-144, pls 1-9. New York. Geological Society of America Bulletin, 46: Clarke, J. M. 1921 (reprinted 1923). James Hall of Albany, 781-868. Geologist and Palaeontologist. Albany, New York, 565 pp, Cooper, G. A., et al. 1942. Correlation of the Devonian sed- pls 1-16. imentary formations of North America. Geological Soci- Clarke, J. M., & D. D. Luther. 1904. Stratigraphic and pa- ety of America Bulletin, 53: 1729-1794. leontologic map of Canandaigua and Naples quadran- Crosse, H., & P.Fischer. 1884. Errata. Journal de Conchyli- gles. New York State Museum Bulletin, 53: 1-67. ologie, 3e série, 32: 415. Clarke, J. M., & R. Ruedemann. 1903. Catalogue of type Denckmann, A. 1903a. Über die untere Grenze des Oberde- specimens of Paleozoic fossils in New York State Mu- von im Lennetal und im Honnetal. Zeitschrift der seum. New York State Museum Bulletin, 65: 1-847. Deutschen Geologischen Gesellschaft, 56: 393-402. Clausen, C.-D. 1969. Oberdevonische Cephalopoden aus Denckmann, A. 1903b. Über Triainoceras costatum A. V. dem Rheinischen Schiefergebirge. II. Gephuroceratidae, Zeitschrit der Deutschen Geologischen Gesellschaft, 55: 85- Beloceratidae. Palaeontographica, Abteilung A, 132: 95- 92. 178, pls 22-26. Denckmann, A. 1906. Über eine Exkursion in das Devon Clausen, C.-D. 1971. Geschichte, Umfang und Evolution und Culmgebeit nordlich von Letmath. Jahrbuch der Gephuroceratidae (Ceph.; Oberdevon.) in heutiger Preussischen Geologischen Landesanstalt, 27: 20-47. 192 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

de Witt, W., Jr. 1960. Java Formation of Late Devonian age Frech, F. 1887. Die palaeozoischen Bildungen von Cabri- in western and central New York. American Association of eres. Zeitschrift deutsch geologischen Gesellschaft, 39: 360- Petroleum Geologists Bulletin, 44: 1933-1939. 487. de Witt, W., Jr., & G. W. Colton. 1959. Revised correlation Frech, F. 1897. Lethaea geognostica. 1 Theil, Lethaea palaeo- of lower Upper Devonian rocks in western and central zoica. 2 Band, 1 Leiferung. Stuttgart, a, b + 256 pp. New York. American Association of Petroleum Geologists Frech, F. 1902a. Über Devonische Ammonoideen. Beiträge Bulletin, 43: 2810-2828. zur Paläontologie und Geologie Österreich-Ungarns und de Witt, W., Jr., & G. W. Colton. 1978. Physical stratigra- des Orients, 14: 27-111. phy of the Genesee Formation (Devonian) in western Frech, F.1902b. Über Epitornoceras und Tornoceras. Zentral- and central New York. United States Geological Survey blatt für Mineralogie, Geologie und Paläontologie, 1902: Professional Paper, 1032-A: 1-22. 172-176. Dreverman, F. 1901. Die Fauna der oberdevonischen Tuff- Frech, F. 1913. Ammoneae Devoniae (Clymenidae, Aphyl- breccie von Langenaubach bei Haiger. Jahrbuch Preussen litidae, Gephuroceratidae, Cheiloceratidae). Fossilium Geologischen Landenanstalt (Berlin), 21: 99-207. Catalogus, Animalia, 42 pp. Ettensohn, F. R. 1985. Controls on development of Catskill Friedman, G. M. 1998. Charles Lyell in New York State. Pp Delta complex basin-facies. Pp 65-77, in: The Catskill 53-69, in: Lyell: the Past is the Key to the Present, D. J. Delta, D. L. Woodrow & W. D. Sevon (eds), Geological Blundell & A. C. Scott (eds), Geological Society of Lon- Society of America Special Paper 201, Boulder, Col- don, Special Publication 143. orado. Glenister, B. F. 1958. Upper Devonian ammonoids from Ettensohn, F. R. 1994. Tectonic control on formation and the Manticoceras Zone, Fitzroy Basin, Western Australia. cyclicity of major Appalachian unconformities and as- Journal of Paleontology, 32: 58-96. sociated stratigraphic sequences. Pp 217-242, in: Tec- Glenister, B. F., D. W. Hanson, & J. E. Preslicka. 2002. tonic and Eustatic Controls on Sedimentary Cycles, J. M. Upper Devonian (Frasnian) ammonoids from the Dennison & F. R. Ettensohn (eds), SEPM (Society for Amana Beds at Price Creek, Iowa. M. A. P.S. Digest, Of- Sedimentary Geology), Concepts in Sedimentology and ficial Publication of the Mid-America Paleontology Society, Paleontology 4, Tulsa, Oklahoma. EXPO XXIV ed., 25(3): 80(2)-86(8), 87-89a (text-figs Feist, R. (ed.). 1990. The Frasnian-Famennian Boundary and 1-6). Adjacent Strata of the Eastern Montagne Noire, France. Grabau, A. W. 1898-1899 (reprinted 1994). Geology and Guidebook, IUGS Subcommission on Devonian Palaeontology of Eighteen Mile Creek and the Lake Shore Stratigraphy, Montpellier, 66 pp. Section of Erie County, New York. Buffalo Society of Nat- Fenton, C. L. 1919. The Hackberry stage of the Upper De- ural Science, Buffalo, New York, xxiv + 402 pp. Reprint vonian. American Journal of Science, 48: 355-376. by Hamburg Natural History Society, Persimmon Press, Fenton, C. L., & M. A. Fenton. 1924. The stratigraphy and Buffalo, New York. fauna of the Hackberry stage of the Upper Devonian. Grasso, T. X., J. W. Harrington, & W.T. Kirchgasser. 1986. Michigan University, Museum of Geology, Contributions, Stratigraphy and paleontology – around Cayuga Lake 1: 1-260. once again. Pp 167-197, in: Field Trip Guidebook, New Fisher, D. W. 1962. Small conoidal shells of uncertain affini- York State Geological Association, 58th Annual Meeting, ties. Pp W98-W143, in: Treatise on Invertebrate Paleon- Cornell University, Ithaca, New York, Hempstead, New tology, Part W, Miscellania, R. C. Moore (ed.), Geological York. Society of America and University of Kansas Press, Grimm, M. 1998. Systematik und Paläoökologie der Bu- Lawrence, Kansas. chiolinae nov. subfamily. Schweizerishe Paläontologische Flower, R. H., & K. E. Caster. 1935. The cephalopod fauna Abhandlungen, 118: 1-176. of the Conewango Series in New York and Pennsylvania. Grünenberg, H. 1925. Beiträge zur Kenntnis des Oberde- Bulletins of American Paleontology, 22(75): 1-74. von der Herzkämper Mulde. Jahresbericht Naturwis- Foord, A. H., & G. C. Crick. 1897. Catalogue of the Fossil senschaftlich Ver Elberfeld, 15: 48-96. Cephalopoda in the British Museum (Natural History), Gürich, G. 1896. Die palaeozoicum im Polnischen Mittel- Part 3, Containing the Bactritidae and Part of the Suborder gebirge. Verhandlungen der Russisch-Kaiserlichen Miner- Ammonoidea. London, 303 pp. alogeischen Gesellschaft zu St. Petersburg, series 2, 32: Fox, I. W. 1932. Geology of part of Finger Lakes region, 1-539. New York. American Association of Petroleum Geologists Hall, J. 1840. Fourth annual report of the survey of the Bulletin, 16: 675-690. fourth geological district. New York Geological Survey An- HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 193

nual Report, 4: 389-456. University of Rochester, Rochester, New York. Hall, J. 1843. Geology of New York. Part IV, Comprising the Hinde, G. J. 1879. On conodonts from the Chazy and Survey of the Fourth Geological District. Carrol & Cook, Cincinnati Group of the Cambro-Silurian, and from the Albany, New York, xxv + 683 pp. Hamilton and Genesee shale divisions of the Devonian, Hall, J. 1860. Notes and observations upon the fossils of the in Canada and the United States. Quarterly Journal of Goniatite limestone in the Marcellus shale of the Hamil- the Geological Society of London, 35: 351-369. ton group in New York and those of the Goniatite beds Holzapfel, E. 1882. Die Goniatiten-Kalke von Adorf in of Rockford, Indiana; with some analogous forms from Waldeck. Palaeontographica, 28: 227-261. the Hamilton group proper. New York State Cabinet of Holzapfel, E. 1899. Die Cephalopoden des Domanik im Natural History Annual Report, 13: 95-112, 125. südlichen Timan. Geologicheskago Komiteta Trudy, 12(3): Hall, J. 1874. Descriptions of New Species of Goniatitidae, 1-56. With a List of Previously Described Species. Albany, New House, M. R. 1961. Acanthoclymenia the supposed earliest York, 4 pp. Devonian clymenid is a Manticoceras. Palaeontology, 3: Hall, J. 1875. Descriptions of new species of Goniatitidae, 472-476. with a list of previously described species. New York State House, M. R. 1962. Observations on the ammonoid succes- Museum, Natural History Annual Report, 27: 132-136. sion of the North American Devonian. Journal of Pale- Hall, J. 1876. Illustrations of Devonian fossils: Gasteropoda, ontology, 3: 247-284. Pteropoda, Cephalopoda, Crustacea and Corals of the House, M. R. 1963. Devonian ammonoid successions and Upper Helderberg, Hamilton and Chemung Groups. New facies in Devon and Cornwall. Quarterly Journal Geolog- York Geological Survey and Weed, Parsons, Albany, New ical Society London, 119: 1-27, pls 1-4. York, 7 pp, 74 pls (Mollusca), 23 pls (Crustacea), 39 pls House, M. R. 1965. A study in the Tornoceratidae; the suc- (corals). cession of Tornoceras and related genera in the North Hall, J. 1879. Descriptions of the Gasteropoda, Pteropoda American Devonian. Transactions of the Royal Society of and Cephalopoda of the Upper Helderberg, Hamilton, London, series B., 250: 79-130. Portage and Chemung Groups. New York Geological Sur- House, M. R. 1968. Devonian ammonoid zonation and vey, Palaeontology of New York, 4(2): xv + 492 pp, 113 correlation between North America and Europe. Pp pls. 1061-1068, in: International Symposium on the Devonian Hall, J. 1886. Fifth Annual Report of the State Geologist for System, Calgary, 1967, vol. 2, D. H. Oswald (ed.), Al- 1885. New York Assembly Document 105, 47 pp., 14 berta Society of Petroleum Geologists, Calgary, Alberta, pls. Canada. Hall, J. 1888. Supplement, containing descriptions and il- House, M. R. 1971. The goniatite wrinkle layer. Smithson- lustrations of Pteropoda, Cephalopoda and Annelida. ian Contributions to Paleobiology, 3: 23-31. New York Geological Survey, Palaeontology of New York, House, M. R. 1975a. Facies and time in Devonian tropical 5(2), supplement: 42 pp. areas. Proceedings of the Yorkshire Geological Society, 40(2, Hall, J., & J. M. Clarke. 1888. Description of the trilobites 16): 233-288. and other Crustacea of the Oriskany, Upper Helderberg, House, M. R. 1975b. Faunas and time in the marine De- Hamilton, Portage, Chemung and Catskill groups. New vonian. Proceedings of the Yorkshire Geological Society, York Geological Survey, Palaeontology of New York, 7: lxiv 40(4, 27): 459-490. + 236 pp, 36 pls House, M. R. 1978. Devonian ammonoids from the Ap- Harris, G. D. 1899. Key to the Upper Devonian of Southern palachians and their bearing on international zonation New York. Elementary Natural History Series, 2. Harris and correlation. Special Papers in Palaeontology, 23: v + Company, Ithaca, New York, 26 pp, 13 pls. 70 pp, 10 pls. Heckel, P. H. 1973. Nature, origin, and significance of the House, M. R. 1979. Biostratigraphy of the early Am- Tully Limestone. Geological Society of America Special monoidea. Special Papers in Palaeontology, 23: 263-280. Paper, 138: 244 pp. House, M. R. 1981a. On the origin, classification and evo- Heckel, P. H. 1997. Overview of the Tully Limestone. Pp lution of the early Ammonoidea. Pp 3-36, in: The Am- 79-100, in: Devonian Cyclicity and Sequence Stratigraphy monoidea, The Evolution, Classification, Mode of Life and in New York State, Field Trip Guide, International Union Geological Usefulness of a Major Fossil Group,M.R. of Geological Sciences, Subcommission on Devonian Stratig- House & J. R. Senior (eds), The Systematics Association raphy, July 22-27, 1997, C. E. Brett & C. A. Ver Straeten Special Volume, number 18, Academic Press, London. (eds), Department of Earth and Environmental Sciences, House, M. R. 1981b. Lower and Middle Devonian goniatite 194 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

biostratigraphy. Pp 33-37, in: Devonian Biostratigraphy of ern Timan (NE Russian Platform). Geological Society Spe- New York, Part 1, Text, W. A. Oliver, Jr., & G. Klapper cial Publication, 178: 147-176. (eds), International Union of Geological Sciences, Sub- House, M. R., & A. E. H. Pedder. 1963. Devonian goni- commission on Devonian Stratigraphy, July 1981, State atites and stratigraphical correlations in Western Canada. University of New York, Binghamton, Washington, DC. Palaeontology, 6: 491-539. House, M. R. 1983. Devonian eustatic events. Proceedings of House, M. R., & J. D. Price. 1985. New late Devonian gen- the Ussher Society, 5: 396-405. era and species of tornoceratid goniatites. Palaeontology, House, M. R. 1985. Correlation of Mid-Paleozoic am- 28: 159-188. monoid evolutionary events with global sedimentary House, M. R., & W. Ziegler. 1977. The goniatite and con- perturbations. Nature, 313: 17-22. odont sequence in the early Upper Devonian at Adorf, House, M. R. 1987. Geographic distribution of Nautilus Germany. Geologica et Palaeontologica, 11: 69-108, pls shells. Pp 53-64, in: Nautilus: the Biology and Paleobiol- 1-5. ogy of a Living Fossil, W. B. Saunders & N. H. Landman House, M. R., & W. Ziegler (eds). 1997. On sea-level fluc- (eds), Plenum Press, New York. tuations in the Devonian. Courier Forschungsinstitut House, M. R. 1995. Devonian precessional and other signa- Senckenberg, 109: 1-146. tures for establishing a Givetian timescale. Pp 37-49, in: Huddle, J. 1974. Middle/Upper Devonian conodont zona- Orbital Forcing Timescales and Cyclostratigraphy, M. R. tion in western New York (abstract). Abstracts with Pro- House (ed.), Geological Society Special Publication 85, grams, North-Central Section, Geological Society of London. America, 6(6): 516. House, M. R. 1996. Juvenile goniatite survival strategies fol- Huddle, J. 1981. Conodonts from the Genesee Formation lowing Devonian extinction events. Geological Society in Western New York. United States Geological Survey Special Publication, 102: 163-185. Professional Paper, 1032-B: 1-66, 31 pls. House, M. R. 2002. Strength, timing, setting, and causes of Hussakof, L., & W. Bryant. 1918. Catalog of the fossil fishes mid-Paleozoic extinctions. Pp 5-25, in: Late Devonian in the Museum of the Buffalo Society of Natural Sci- Biotic Crisis: Ecological, Depositional and Geochemical ences. Buffalo Society of Natural Sciences Bulletin, 12: 1- Records, G. Racki & M. R. House (eds). Palaeogeogra- 346. phy, Palaeoclimatology, Palaeoecology, 181. Hyatt, A. 1884. Genera of fossil cephalopods. Boston Society House, M. R., R. T. Becker, R. Feist, G. Flajs, C. Girard, & of Natural History Proceedings, 22: 253-338. G. Klapper. 2000. The Frasnian/Famennian boundary Hyatt, A. 1900. Class 5, Cephalopoda. Pp 502-604, in: GSSP at Coumiac, Southern France. Courier Textbook of Palaeontology, vol. 1, K. A. Zittel & C. R. Forschungsinstitut Senckenberg, 225: 59-75. Eastman (eds), London. House, M. R., R. Feist, & D. Korn. 2000. The International Commission on Zoological Nomenclature Middle/Upper Devonian boundary GSSP at Puech de [ICZN]. 1956. Opinion 392. Emendation to Sandberg- la Suque, Southern France. Courier Forschungsinstitut eroceras of the generic name Sandbergeoceras Hyatt, 1844 Senckenberg, 225: 49-58. (Class Cephalopoda, Order Ammonoidea). Opinions House, M. R., M. Gordon, Jr., & W. J. Hlavin. 1986. Late and Declarations Rendered by the International Commis- Devonian ammonoids from Ohio and adjacent states. sion on Zoological Nomenclature, 3(15): 295-304. Journal of Paleontology, 60: 126-144. International Commission on Zoological Nomeclature House, M. R., and W. T. Kirchgasser. 1993. Devonian go- [ICZN]. 1999. International Code of Zoological Nomen- niatite biostratigraphy and timing of facies movements clature, 4th ed. International Trust for Zoological in the Frasnian of eastern North America. Pp 267-292, Nomenclature, London, 306 pp. in: High Resolution Stratigraphy, E. A. Hailwood & R. Johnson, J. G. 1970. Taghanic Onlap and the end of North B. Kidd (eds), Geological Society Special Publication 70, American Devonian provinciality. Geological Society of London. America Bulletin, 81: 2077-2106. House, M. R., W. T. Kirchgasser, J. D. Price, & G. Wade. Johnson, J. G. 1974. Extinction of perched faunas. Geology, 1985. Goniatites from the Frasnian (Upper Devonian) 2: 479-482. and adjacent strata of the Montagne Noire. Hercynica, Johnson, J. G. 1989. Base of the Upper Devonian in the 1: 1-21. conodont zonation. Newsletters in Stratigraphy, 21: 11- House, M. R., V. V. Menner, R. T. Becker, G. Klapper, N. 145. S. Ovnatanova, & V. Kuz'min. 2000. Reef episodes, Johnson, J. G., G. Klapper, & C. A. Sandberg. 1985. De- anoxia and sea level changes in the Frasnian of the south- vonian eustatic fluctuations in Euramerica. Geological HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 195

Society of America Bulletin, 96: 567-587. Kirchgasser, W. T. 1974. Notes on the ammonoid and con- Johnson, J. G., G. Klapper, & C. A. Sandberg. 1986. Late odont zonations of southwestern New York. Pp B9-B13, Devonian eustatic cycles around margin of Old Red in: Field Trip Guidebook, New York State Geological Asso- Sandstone Continent. Annales de la Société Géologique de ciation, 46th Annual Meeting, D. N. Peterson (ed.), State Belgique, 103: 141-147. University of New York, Fredonia, New York. Johnson, K. G., & G. M. Friedman. 1969. The Tully clastic Kirchgasser, W. T. 1975. Revision of Probeloceras Clarke, correlatives (Upper Devonian) of New York State: a 1898 and related ammonoids from the Upper Devonian model for recognition of alluvial dune (?), tidal, of western New York. Journal of Paleontolology, 49: 58- nearshore (bar and lagoon), and offshore sedimentary 90, pls 1-3. environments in a tectonic delta complex. Journal of Sed- Kirchgasser, W. T. 1981. Stop 1B, Eighteen Mile Creek at imentary Petrology, 39: 451-485. RR, and Stop 13A, B Mill Creek Glen, Lodi. Pp 9-11, Keyserling, A. 1844. Beschreibung einiger Goniatiten aus 44-45, in: Devonian Biostratigraphy of New York, Part 2, dem Domanik-Schiefer. Verhandlungen der kaiserlich-rus- Stop Descriptions, W. A. Oliver, Jr., & G. Klapper (eds), sischen Mineralogischen Gesellschaft, 1844: 217-238. International Union of Geological Sciences, Subcom- Keyserling, A. 1846. Wissenschaftliche Beobachtungen auf mission on Devonian Stratgraphy, July 1981, State Uni- einer Reise in das Petschora-Land im Jahre 1843. St. Pe- vesity of New York, Binghamton, Washington, DC. tersburg, 467 pp. Kirchgasser, W. T. 1982. Ammonoid correlation near the Kilfoyle, C. F. 1969. Catalog of type specimens in the New delta front: Late Devonian Genesee Formation, New York State Museum, supplement 6. New York State Mu- York (abstract). Northeastern/Southeastern Sections Geo- seum and Science Service Bulletin, 413, 305 pp. logical Society of America, Abstracts with Programs, 14(1- Kindle, E. M. 1896. The relation of the Ithaca Group to the 2): 31. faunas of the Portage and Chemung. Bulletins of Ameri- Kirchgasser, W.T. 1985. Ammonoid horizons in the Upper can Paleontology, 2(6): 56 pp. Devonian Genesee Formation of New York: legacy of Kindle, E. M. 1901. The Devonian fossils and stratigraphy Genesee, Portage and Chemung. Pp 225-235, in: The of Indiana. Indiana Department of Geology and Natural Catskill Delta, D. L. Woodrow & W. D. Sevon (eds), Resources, Annual Report, 25(for 1900): 529-758. Geological Society of America Special Paper 201, Boul- Kirchgasser, W. T. 1965. The Parrish Limestone (Upper De- der, Colorado. vonian) of West Central New York. Unpublished M. S. Kirchgasser, W. T. 1994. Early morphotypes of Ancyrodella Thesis, Cornell University, Ithaca, New York, 177 pp, 5 rotundiloba at the Middle/Upper Devonian boundary, pls. Genesee Formation, west-central New York. Pp 117- Kirchgasser, W.T. 1967. Paleontology and Stratigraphy of the 134, in: Studies in Stratigraphy and Paleontology in Honor Concretions and Limestones of the Upper Devonian of Donald W. Fisher, E. Landing (ed.), New York State Cashaqua Shale Member, Sonyea Formation, New York. Museum Bulletin 481, Albany, New York. Unpublished Ph. D. Dissertation, Cornell University, Kirchgasser, W.T. 1996a. Evidence of distant transport (bas- Ithaca, New York, 82 pp, 9 pls. inward) of reworked conodonts in a condensed sequence Kirchgasser, W. T. 1968. Paleontology and stratigraphy of in the Upper Devonian (lower Frasnian) of western New the concretions and limestones of the Upper Devonian York (abstract). Abstracts, 30th International Geological Cashaqua Shale Member, Sonyea Formation, New York Congress, Beijing, China, 2: 86. (abstract). Dissertation Abstracts, 28B: 4175-4176. Kirchgasser, W. T. 1996b. Correlation of ammonoid-con- Kirchgasser, W. T. 1969. Stratigraphic relations within the odont event-beds in the late Devonian of the Northern Frasnian Cashaqua shale member (Sonyea Formation) Appalachians, North America (abstract). Abstract with of New York (abstract). Abstracts with Programs for 1969, Program, International Commission on Stratigraphy, Sub- Part 1, Northeastern Section, Geological Society of America: commission on Devonian Stratigraphy Meeting, August 8, 33-34. 1996. 30th International Geological Congress, Beijing: 4. Kirchgasser, W. T. 1973. Lower Upper Devonian stratigra- Kirchgasser, W. T. 1998. Problems in sampling the North phy from the Batavia Warsaw meridian to the Genesee Evans and Genundewa Limestones (Genesee Group of valley: goniatite sequence and correlation. Pp C1-C21, New York) and the development of the conodont zona- in: Guidebook to Field Trips, New York State Geological tion around the Middle-Upper Devonian (Givetian- Association, 45th Annual Meeting, P. C. Hewitt (ed.), Frasnian) boundary (abstract). Abstracts with Programs, State University of New York, Brockport, and Monroe North Central Section, Geological Society of America, Community College, Brockport, New York. 30(2): 27. 196 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

Kirchgasser, W.T. 2001. Taphonomy and “sequence” of con- G. Klapper (eds), International Union of Geological Sci- odonts and ichthyoliths in the North Evans remanié de- ences, Subcommission on Devonian Stratigraphy, July posit at the Taghanic-sub-Genundewa Unconformity 1981, State University of New York, Binghamton, Wash- (late Givetian-early Frasnian) in western New York. P. ington, DC. 53, in: Mid-Paleozoic Bio- and Geodynamics, The North Kirchgasser, W. T., & D. Koslowski. 1996. North Evans Gondwana-Laurussia Interaction, 15th International conodont fauna at Cayuga Creek, Erie County, western Senckenberg Conference, P. Königshof, G. Plodowski, & New York: evidence of reworked conodonts during part E. Schindler (eds), Senckenbergische Naturforschende of the Early Late Devonian (abstract). Abstracts with Pro- Gesellschaft, Frankfurt am Main. grams, Northeastern Section, Geological Society of America, Kirchgasser, W.T. 2002. Taphonomy of conodonts and mi- 28(3): 72-73. crovertebrates in remanié horizons: new approaches to Kirchgasser, W.T., W. A. Oliver, Jr., & L. V. Rickard. 1986. unraveling stratigraphic relations around the Middle- Devonian series boundaries in the eastern United States. Upper (Givetian-Frasnian) Devonian boundary in west- Pp 233-260, in: Devonian Series Boundaries – Results of ern New York (abstract). Abstracts with Programs, World-Wide Studies, W. Ziegler & R. Werner (eds), Northeastern Section, Geological Society of America, 34(1): Courier Forschungsinstitut Senckenberg, 75. 58. Kirchgasser, W. T., D. J. Over, & D. L. Woodrow. 1994. Kirchgasser, W.T. 2004. Conodonts in pyrite lag deposits at Frasnian (Upper Devonian) strata of the Genesee River the Taghanic Unconformity in New York state: problems Valley, western New York State. Pp 325-358, in: Field in dating faunas in highly condensed beds around the Trip Guidebook, New York State Geological Association, Middle (Givetian)-Upper (Frasnian) Devonian bound- 66th Annual Meeting, C. E. Brett & J. Scatterday (eds), ary (abstract). Devonian Neritic-Pelagic Correlation and Department of Earth and Environmental Sciences, Uni- Events, Abstracts, IUGS Subcommission on Devonian versity of Rochester, Rochester, New York. Stratigraphy and Institut Scientifique, University Mo- Kirchgasser, W. T., D. J. Over, & D. L. Woodrow. 1997. hammed V, Rabat, Morocco: 27. Upper Devonian facies, strata and cycles in western New Kirchgasser, W. T., C. E. Baird, & G. C. Brett. 1989. Re- York State. Pp 131-146, 232, in: Devonian Cyclicity and gional placement of Middle/Upper Devonian (Givetian- Sequence Stratigraphy in New York State, Field Trip Guide- Frasnian) boundary in western New York State. Pp book, International Union of Geological Sciences, Subcom- 113-117, in: The Devonian of the World, Proceedings of mission on Devonian Stratigraphy, July 22-27, 1997, C. the Second International Symposium on the Devonian Sys- E. Brett & C. A. Ver Straeten (eds), Department of tem, Calgary, Canada, vol. 3, N. J. McMillan, A. F. Earth and Environmental Sciences, University of Embry, & D. J. Glass (eds), Canadian Society of Petro- Rochester, Rochester, New York. leum Geologists Memoir 14, Calgary, Alberta, Canada. Kirchgasser, W. T., & B. Vargo. 1998. Middle Devonian Kirchgasser, W. T., & C. E. Brett. 1981. Stop 5, Cayuga conodonts and ichthyoliths in an Upper Devonian lime- Creek. Pp 18-20, in: Devonian Biostratigraphy of New stone in New York: implications for correlations around York, Part 2, Stop Descriptions, W. A. Oliver, Jr., & G. the Givetian-Frasnian boundary (abstract). P.82, in: Ab- Klapper (eds), International Union of Geological Sci- stracts, Seventh European Conodont Symposium (ECOS ences, Subcommission on Devonian Stratgraphy, July VII), G. Bagnoli (ed.), Tipografia Compositori Bologna, 1981, State Univesity of New York, Binghamton, Wash- Bologna-Modena. ington, DC. Klapper, G. 1981. Review of New York Devonian conodont Kirchgasser, W. T., C. E. Brett, & G. C. Baird. 1997. Se- biostratigraphy. Pp 57-66, in: Devonian Biostratigraphy of quences, cycles and events in the Devonian of New York New York, Part 1, Text, W. A. Oliver, Jr., & G. Klapper State: an update and overview. Pp 5-21, 194-198, in: (eds), International Union of Geological Sciences, Sub- Devonian Cyclicity and Sequence Stratigraphy in New York commission on Devonian Stratigraphy, July 1981, State State, Field Trip Guidebook, International Union of Geo- University of New York, Binghamton, Washington, DC. logical Sciences, Subcommission on Devonian Stratigraphy, Klapper, G. 1988. Intent and reality in biostratigraphic July 22-27, 1997, C. E. Brett & C. A. Ver Straeten (eds), zonation; a reply to Sandberg, Ziegler and Bultynck. Department of Earth and Environmental Sciences, Uni- Newsletters in Stratigraphy, 19: 179-183. versity of Rochester, Rochester, New York. Klapper, G. 1989. The Montagne Noire Frasnian (Upper Kirchgasser, W.T., & M. R. House. 1981. Upper Devonian Devonian) conodont succession. Pp 449-468, in: De- goniatite biostratigraphy. Pp 39-55, in: Devonian Bios- vonian of the World, Proceedings of the Second Interna- tratigraphy of New York, Part 1, Text, W. A. Oliver, Jr., & tional Symposium on the Devonian System, Calgary, HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 197

Canada, vol. 3, N. J. McMillan, A. F. Embry, & D. J. Members of the Genesee Formation, Western New York Glass (eds), Canadian Society of Petroleum Geologists State. Unpublished M. S. Thesis, University of Iowa, Memoir 14, Calgary, Alberta, Canada. Iowa City, Iowa, 57 pp. Klapper, G. 1997. Graphic correlation of the Frasnian Kralick, J. A. 1994. The conodont genus Ancyrodella in the (Upper Devonian) sequences of the Montagne Noire, Middle Genesee Formation (Lower Upper Devonian, France and western Canada. Pp 113-129, in: Paleozoic Frasnian), western New York. Journal of Paleontology, 68: Sequence Stratigraphy, Biostratigraphy, and Biogeography: 1384-1395, figs 3-6. Studies in Honor of J. G. ("Jess") Johnson, G. Klapper, M. Krebs, W. 1979. Devonian basinal facies. Special Papers in A. Murphy, & J. A. Talent (eds), Geological Society of Palaeontology, 23: 125-139. America Special Paper 321, Boulder, Colorado. Kulicki, C., K. Tanabe, N. H. Landman, & R. H. Mapes. Klapper, G. 2000. Species of Spathognathodontidae and 2001. Dorsal shell wall in ammmonoids. Acta Palaeon- Polygnathidae (Conodonta) in the recognition of Upper tologica Polonica, 46: 23-42. Devonian stage boundaries. Courier Forschungsinstitut Kullmann, J., & J. Wiedmann. 1970. Significance of sutures Senckenberg, 220: 153-159. in phylogeny of ammonoids. University of Kansas Paleon- Klapper, G., & R. T. Becker. 1999 (1998). Comparison of tological Contributions, 47: 1-32. Frasnian (Upper Devonian) conodont zonations. Pp Kullmann, J., & W. Ziegler. 1970. Conodonten und Goni- 339-348, in: Studies on Conodonts - Proceedings of the Sev- atiten von der Grenze Mittle /Oberdevon aus dem Profil enth European Conodont Symposium, E. Serpagli (ed.), am Martenberg (Ostrand des Rheinischen Schieferge- Bollettino della Società Paleontologica Italiana, 37(2-3). birges). Geologica et Palaeontologica, 4: 73-85, pl. 1. Klapper, G., R. Feist, R. T. Becker, & M. R. House. 1993. Landing, E., & C. E. Brett. 1985. Anatomy of a disconfor- Definition of the Frasnian/Famennian Stage boundary. mity: sedimentology and trace fossils of the basal Hun- Episodes, 16: 433-441. gry Hollow Formation (Middle Devonian, Hamilton Klapper, G., R. Feist, & M. R. House. 1987. Decision on group, southwestern Ontario) (abstract). Abstracts with the boundary for the Middle/Upper Devonian Series Programs, Geological Society of America, 17(1): 30. Boundary. Episodes, 10: 97-101. Landing, E., & C. E. Brett. 1987. Trace fossils and regional Klapper, G., & J. G. Johnson. 1990. Revisions of Middle significance of a Middle Devonian (Givetian) disconfor- Devonian conodont zones. Journal of Paleontology, 64: mity in southwestern Ontario. Journal of Paleontology, 934-936, 941. 61(2): 205-230. Klapper, G., W. T. Kirchgasser, & J. F. Baesmann. 1995. Landman, N. H., K. Tanabe, & Y. Shigeta. 1995. Am- Graphic correlation of a Frasnian (Upper Devonian) monoid embryonic development. Pp 343-405, in: Am- composite standard. Pp 177-184, in: Graphic Correla- monoid Paleobiology, N. H. Landman, K. Tanabe, & R. tion, K. O. Mann & H. R. Lane (eds), SEPM (Society A. Davis (eds), Plenum Press, New York and London. for Sedimentary Geology), Special Publication 53, Tulsa, Lincoln, D. F.1895. Report on the structural and economic Oklahoma. geology of Seneca County. New York State Museum, 48th Klapper, G., G. M. Philip, & J. H. Jackson. 1970. Revision Annual Report of the Regents, 1894, 2: 60-125. of the Polygnathus varcas group (Conodonta, Middle De- Lindemann, R. H. 2002. Dacryoconarid bioevents of the vonian). Neues Jahrbuch für Geologie und Paläontologie, Onondaga Formation and Marcellus Subgroup, Cherry Monatshefte, 11: 650-667. Valley, New York. Pp B7-1-B7-17, in: Guidebook for Kloc, G. J. 1983. Stratigraphic Distribution of Ammonoids Fieldtrips in New York and Vermont, 94th Annual Meet- from the Middle Devonian Ludlowville Formation in New ing, New England Geological Conference and 74th Annual York. Unpublished M. A. Thesis, University of Buffalo, Meeting, New York State Geological Association, Lake Buffalo, New York, 78 pp. George, New York, J. M. McLelland & P.Karabinos (eds), Korn, D. 1986. Ammonoid evolution in Late Famennian University of Maine, Orono, Maine. and Early Tournasian. Annales Société Geologigue de Bel- Loomis, F. B. 1903. The dwarf fauna of the pyrite layer at gique, 109: 49-54. the horizon of the Tully Limestone in western New York. Korn, D. 1991. Threedimensionally preserved clymenids New York State Museum Bulletin, 69: 892-920. from the Hangenberg Black Shale of Drewer. Neues Luppold, F. W., C. D. Clausen, D. Korn, & D. Stoppel. Jahrbuch für Geologie und Paläontologie, Monatsheft, 9: 1994. Devon/Karbon-Grenzprofile in Berich von Rem- 553-563. scheid-Altenaer Sattle, Warsteiner Sattle, Briloner Sattel Kralick, J. A. 1991. Conodont Biostratigraphy Within the und Attendorn-Elsper Doppelmulde (Rheinisches Upper Devonian (Frasnian) Penn Yan and Genundewa Schiefergebirge). Pp 7-69, in: Die Devon/Karbon-Grenze 198 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

im Rheinischen Schiefergebirge, A. Hendricks (ed.), Ge- Triasprovinz. Abhandlungen Kaiserkönig Geologischen Re- ologie und Palaöntologie in Westfalen 29, Westfälsiches ichsanstalt, 10: 1-322, pls 1-94. Museum für Naturkunde, Münster. Montesinos, J. R. 1987. Aulatornoceras (Truyolsoceras) n. Luther, D. 1903. Stratigraphy of the Portage Formation be- subgen. (Ammonoidea, Tornoceratina) del Devonico de tween the Genesee Valley and Lake Erie. New York State las cordilleras Cantabrica e Iberica (NO y NE de Es- Museum Bulletin, 69: 1000-1029. pana). Cuadernos Labatorio Xeologio de Laxe Coruna, 12: Luther, D. 1910 (1909). Nun-da-wa’-o, the oldest Seneca 119-126. village. New York State Museum Bulletin, 140(6th Report Montesinos, J. R., & A. H. Henn. 1986. La fauna de Phar- of the Director): 213-221. ciceras (Ammonoidea) de la Formación Cardaño (Do- Luther, D. 1911. Geology of the Honeoye-Wayland Quad- minio Palentino, Cordillera Cantabrica, NO de España. rangles. New York State Museum Bulletin, 152: 5-29. Trabajos de Geologia, 16: 61-76. Lyashenko [Ljaschenko], G. P. 1957. Novye vidy Devon- Moore, R. C. (ed.). 1957. Treatise on Invertebrate Paleontol- skikh goniatitov [New species of Devonian goniatites]. ogy, Part L, Mollusca 4, Cephalopoda, Ammonoidea. Ge- Trudy Vsesoiuznogo Nftianogog Nauchnoissledovatel’skogo ological Society of America and University of Kansas Geolgorazvedochnogo Instiuto, 8: 192-211. Press, Lawrence, Kansas, L490 pp. MacDonald, K. B. 1975. Quantitative community analysis: Müller, K. J. 1956. Cephalopodenfauna und Stratigraphie recurrent and cluster techniques applied to the fauna of des Oberdevons von Schleiz und Zeulenroda in Thürin- the Upper Devonian Sonyea Group, New York. Journal gen. Geologisches Jahrbuch, 20: 156-196. of Geology, 82: 473-499. Münster, G., Graf zu. 1843. Nachtraf zu den Versteinerun- Maeda, H., & A. Seilacher. 1995. Ammonoid Taphonomy. gen des Übergangskalkes mit Clymenien von Ober- Pp 543-578, in: Ammonoid Paleobiology, N. H. Land- franken. Beiträge Petrefacten-Kunde (Bayreuth), 5: man, K. Tanabe, & R. A. Davis (eds), Plenum Press, 112-128. New York and London Murphy, A. E., B. B. Sageman, C. A. Ver Straeten, & D. J. Matern, H. 1929. Die Gleiderung der Adorf-Stufe, Zugleich Hollander. 2000. Organic carbon burial and faunal dy- ein Beitrag zur Nomenklatur von Gepuroceras Hyatt namics in the Appalachian Basin during the Devonian (Ceph.). Senckenbergiana, 11: 142-152. (Givetian-Famennian) greenhouse; an integrated pale- Matern, H. 1931a. Das Oberdevon der Dill-Mulde. Ab- oecological and biogeochemical approach. Pp 351-385, handlungen Preussen Geologischen Landesanstalt, Neue in: Warm Climates in Earth History, B. Huber, K. Folge, 134: 1-139, pls 1-4. MacLeod, & S. L. Wing (eds), University of Cambridge Matern, H. 1931b. Die Goniatiten Fauna der Schistes de Press, Cambridge, UK. Matagne in Belgien. Bulletin du Musée Royale d’Histoire Oliver, W. A., Jr., W. de Witt, Jr., J. M. Dennison, D. M. Naturelle de Belgique, 7: 1-15. Hoskins, & J. W. Huddle. 1968. Devonian of the Ap- Miller, A. K. 1938. Devonian ammonoids of America. Ge- palachian basin, United States. Pp 1001-1040, in: Inter- ological Society of America Special Paper, 14: xiii + 202 national Symposium on the Devonian System, Calgary, pp, pls 1-39. 1967, vol. 1, D. H. Oswald (ed.), Alberta Society of Pe- Miller, A. K. 1952. Proposed acceptance of the emendation troleum Geologists, Calgary, Alberta, Canada. to “Sandbergeroceras” of the generic name “Sandbergeo- Oliver, W. A., Jr., W. de Witt, Jr., J. M. Dennison, D. M. ceras” Hyatt, 1884 (Class Cephalopoda, Order Am- Hoskins, & J. W. Huddle. 1969. Correlation of Devon- monoidea); and the establishment of “Goniatites ian rock units in the Appalachian basin. United States tuberculosocostatus” G. and F. Sandberger, 1850, as its Geological Survey Oil and Gas Investigations Chart OC type species. Bulletin of Zoological Nomenclature, 6: 357- 64. 359. Oliver, W. A., Jr., & G. Klapper (eds). 1981a. Devonian Miller, A. K., & R. H. Flower. 1936. A Sporadoceras from Biostratigraphy of New York, Part 1, Text. International America. Journal of Geology, 44: 751-757. Union of Geological Sciences, Subcommission on De- Miller, A. K., & W. M. Furnish. 1957. Anarcestina and Go- vonian Stratigraphy, July 1981, State University of New niatitina. Pp L29-L36, L47-L69, in: Treatise on Inverte- York, Binghamton, Washington, DC, 105 pp. brate Paleontology, Part L, Mollusca 4, Cephalopoda, Oliver, W. A., Jr., & G. Klapper (eds). 1981b. Devonian Ammonoidea, R. C. Moore (ed.), Geological Society of Biostratigraphy of New York, Part 2, Stop Descriptions. In- America and University of Kansas Press, Lawrence, ternational Union of Geological Sciences, Subcommis- Kansas. sion on Devonian Stratigraphy, July 1981, State Mojsisovics, E. 1882. Die Cephalopoden der mediterranen University of New York, Binghamton, Washington, DC, HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 199

69 pp. bung von Balve in Westfalen. Jahrbuch Preussischen Ge- Oppenheimer, J. 1916. Das Oberdevon von Brünn. Ver- ologischen Landesanstalt, 44: 51-97, pls 4-5. handlungen des Naturwissenschaftlichen Vereins in Brünn, Paproth, E., R. Feist, & G. Flajs. 1991. Decision on the De- 54: 156-196. vonian-Carboniferous boundary stratotype. Episodes, Orlov, Iu. A. (ed.). 1962. Osnovy Paleontologii, 5, Molluski- 14(4): 331-336. Golovonogie, 1. Akademiya Nauk SSSR (Moscow), 438 Pepper, J. F., S. Clark, & W. de Witt, Jr. 1985. Nodules of pp, 32 pls. diagenetic barite in Upper Devonian Shales of western Over, D. J. 1997a. Frasnian-Famennian boundary in west- New York. United States Geological Survey Bulletin, 1653: ern New York: Dunkirk Beach and Buffalo Creek Valley. 1-11. Pp 147-155, in: Devonian Cyclicity and Sequence Stratig- Pepper, J. F., & W. de Witt, Jr. 1950. Stratigraphy of the raphy in New York State, Field Trip Guidebook, Interna- Upper Devonian Wiscoy Sandstone and equivalent tional Union of Geological Sciences, Subcommission on Hanover Shale in western and central New York. United Devonian Stratigraphy, July 22-27, 1997, C. E. Brett & States Geological Survey Oil and Gas Investigations Chart C. A. Ver Straeten (eds), Department of Earth and En- OC-37. vironmental Sciences, University of Rochester, Pepper, J. F., & W. de Witt, Jr. 1951. Stratigraphy of the Rochester, New York. late Devonian Perrysburg Formation in western and cen- Over, D. J. 1997b. Conodont biostratigraphy of the Java tral New York. United States Geological Survey Oil and Formation (Upper Devonian) and the Frasnian/ Fa- Gas Investigations Chart OC-45. mennian boundary in New York State. Pp 161-177, in: Pepper, J. F., W. de Witt, Jr., & G. W. Colton. 1956. Stratig- Paleozoic Sequence Stratigraphy, Biostratigraphy, and Bio- raphy of the West Falls Formation of Late Devonian age geography: Studies in Honor of J. G. ("Jess") Johnson, G. in western and central New York. United States Geological Klapper, M. A. Murphy, & J. A. Talent (eds), Geological Survey Oil and Gas Investigations Chart OC-55. Society of America Special Paper 321, Boulder, Col- Petter, G. 1959. Goniatites Devoniennes du Sahara. Publi- orado. cations de la Service de la Carte Geologique de l'Algerie, Over, D. J. 2002. The Frasnian-Famennian boundary in the nouvelle série, Paléontologie, Mémoire, 2: 1-313, pls 1-26. Appalachian Basin, Michigan Basin, Illinois Basin and Quenstedt, F. A. 1846. Petrefaktenkunde Deutschland, Band southern continental margin, central and eastern United 1, Cephalopoden. Tübingen, 580 pp, 36 pls. States. Palaeogeography, Palaeoclimatology, Paleoecology, Rabien, A. 1956. Zur Stratigraphie und Fazies des Oberde- 181: 153-169. vons in der Waldecker Hauptmulde. Abhandlungen des Over, D. J., G. C. Baird, & W. K. Kirchgasser. 1999. Frasn- Hessischen Landesamt für Bodenforschung, 16: 1-83, pls ian (lower Upper Devonian) geology of western New 1-3. York as seen along Eighteenmile Creek and Route 20A: Racki, G., & P.Bultynck. 1993. Conodont biostratigraphy submarine discontinuities, gravity flows and mass extinc- of the Middle to Upper Devonian boundary beds in the tion. Pp B8-B21, in: Field Trip Guidebook, New York Kielce area of the Holy Cross Mts. Acta Geologica State Geological Association, 71st Annual Meeting, G. G. Polonica, 43: 1-25, pls 1-9. Lash (ed.), Department of Geosciences, State University Racki, G., & T. Wrzolek. 1989. Middle-Upper Devonian of New York, Fredonia. boundary: ambiguous reality of its stratotype. Pp 231- Over, D. J., C. A. Conaway, D. J. Katz, W. D. Goodfellow, 236, in: Contributions to Devonian Paleontology and & D. C. Grogoire. 1997. Platinum group element en- Stratigraphy, Part II, Various Devonian Topics, O. H. Wal- richments and possible chondritic Ru:Ir across the Frasn- liser & W. Ziegler (eds), Courier Forschungsinstitut ian-Famennian boundary, western New York state. Senckenberg, 110. Palaeogeography, Palaeoclimatology, Palaeoecology, 132: Richter, R. 1929. Nomenclatur und Nomenklatur. Senck- 399-410. enbergiana, 11: 380-386. Over, D. J., T. H. Hopkins, A. Brill, & A. L. Spaziani. 2003. Rickard, L. V. 1964. Correlation of the Devonian rocks in Age of the Middlesex Shale (Upper Devonian, Frasnian) New York. New York State Museum and Science Service, in New York. Pp 217-223, in: Mid-Paleozoic Bio- and Geological Survey, Map and Chart Series, 3. Geodynamics: the North Gondwana-Laurussia Interaction, Rickard, L. V. 1975. Correlation of the Silurian and Devon- P. Königshof & E. Schindler (eds), Proceedings of the ian rocks in New York state. New York State Museum and 15th International Senckenberg Conference, Courier Science Service, Map and Chart Series, 15. Forschungsinstitut Senckenberg, 242. Rickard, L. V. 1981. The Devonian system in New York. Pp Paekelmann, W. 1924. Das Devon und Karbon der Umbge- 5-22, in: Devonian Biostratigraphy of New York, Part 1, 200 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

Text, W. A. Oliver, Jr., & G. Klapper (eds), International dewa Limestone of Western New York. Unpublished M. S. Union of Geological Sciences, Subcommission on De- Thesis, University of Rochester, Rochester, New York, vonian Stratigraphy, July 1981, State University of New 113 pp. York, Binghamton, Washington, DC. Sayn, G. 1901. Les Ammonites priteuses des marnes Valang- Rickard, L. V. 1984. Correlation of the subsurface Lower iennes du Sud-Est de la France. Mémoires de la Société and Middle Devonian of the Lake Erie region. Geological Géologique de France, 23: 1-27. Society of America Bulletin, 95: 814-828. Schindewolf, O. H. 1922. Eininge Randbermerkungen zu Rickard, L.V., & D. W. Fisher. 1970. Geologic map of New E. Perna’s Abhandlung ‘Die Ammoneen des oberen York. New York State Museum and Science Service, Map Neodevon von Ostabhang des Südurals.’ Senckenber- and Chart Series, 15. giana, 4: 185-196. Roemer, F.1846. Über Devon Formation in Nord Amerika. Schindewolf, O. H. 1923. Beitrage zur Kenntnis der Palaeo- Neues Jahrbuch, 1846: 58-59. zoicums in Oberfranken. Ostthüringen und dem Säch- Rudwick, M. J. S. 1985. The Great Devonian Controversy: sischen Vogtlande. 1. Stratigraphie und Ammoneen- The Shaping of Scientific Knowledge among Gentlemanly fauna des Oberdevons von Hof a. d. Saale. Neues Specialists. University of Chicago Press, Chicago, Illinois, Jahrbuch für Mineralogie, Geologie und Paläontologie, 49: 494 pp. 250-358, 393-509, pls 14-18. Ruzhentsev, V. E. 1957. Filogeneticheskaya sistema paleo- Schindewolf, O. H. 1934. Über eine oberdevonische Am- zoyskikh ammonoidey. Byulleten Moskovskogo Obshch- moneen-Fauna der Rocky-Mountains. Neues Jahrbuch estva Ispytatelei Prirody, Otdel geologicheskii, 32: 49-64. für Mineralogie, Geologie und Paläontologie, 72: 331-350. Ruzhentsev, V. E. 1960. Printsipy sistematiki sistema i filo- Schindewolf, O. H. 1936. Neue Gattungen tiefoberdevonis- geniya Paleozoyskikh ammonoidey. Trudy Paleontologich- ches Goniatiten. Zeitschrift der Deutschen Geologischen eskogo Instituta, Akademiya Nauk SSSR, 83: 1-332. Gesellschaft, 88: 689-691. Ruzhentsev, V. E. (ed.). 1962. Osnovy Paleontologii, 5, Schindewolf, O. H. 1937a. Zwei neue bemerkenswerte Go- Molluski Golovonogie, 1. Izdatel'stvo Akademia Nauk niatiten-Gattungen des rheinischen Oberdevons. SSSR (Moscow), 425 pp. Jahrbuch Preussischen Geologischen Landesanstalt, 58: Sandberg, C. A., J. R. Morrow, & W. Ziegler. 2002. Late 242-256, pl. 1. Devonian sea-level changes, catastrophic events and mass Schindewolf, O. H. 1937b. Stratigraphie und Palaeontolo- extinctions. Pp 473-487, in: Catastrophic Events and gie der Wocklumer Schichten. Abhandlungen der Mass Extinctions: Impacts and Beyond, C. Koeberl & K. Preussischen Geologischen Landesanstalt, Neue Folge, 178: G. MacLeod (eds), Geological Society of America Spe- 1-132, pls 1 4. cial Paper 356, Boulder, Colorado. Schindewolf, O. H. 1940. Konvergenzen bei Korallen und Sandberg, C. A., W. Ziegler, & P.Bultynck. 1988. Middle- bei Ammoneen. Fortschritte in der Geologie und Palaeon- Upper Devonian boundary as an example of intent and tologie, 12: i-vii, 389-492. reality in biostratigraphic zonation. Newsletters on Stratig- Schindewolf, O. H. 1949. Zur Phylogenie der Clymenien raphy, 18: 117-121. (Cephalop., Ammon.). Neues Jahrbuch für Mineralogie, Sandberg, C. A., W. Ziegler, & P. Bultynck. 1989. New Geologie und Paläontologie, Monatshefte, Abteilung B, 7: standard conodont zones and early Ancyrodella phy- 197-209. logeny across Middle-Upper Devonian boundary. Pp Schindewolf, O. H. 1952. Über das Oberdevon und Un- 195-230, in: Contributions to Devonian Paleontology and terkarbon von Saalfeld in Ostthüringen. Senckenber- Stratigraphy, Part II, Various Devonian Topics, O. H. Wal- giana, 32: 281-306, pls 1-3. liser & W. Ziegler (eds), Courier Forschungsinstitut Schindewolf, O. H. 1954. On the development, evolution, Senckenberg, 110. and terminology of ammonoid suture line. Bulletin of Sandberger, G., & F. Sandberger. 1849-1856. Systematische the Museum of Comparative Zoology, 112: 217-237. Beschreibung und Abbildung der Versteinerungen des Schindewolf, O. H. 1955. Zur Taxionomie und Nomen- Rheinischen Schichtensystems in Nassau. Wiesbaden, 564 klatur der Clymenien. Ein Epilog. Neues Jahrbuch für pp, 39 pls. Geologie und Paläontologie, Monatshefte, 10: 417-429. Sartenaer, P. 2003. Tullypothyridina, a new late Givetian Schindler, E. 1990. Die Kellwasser-Krise (hohe Frasnian rhynchonellid (brachiopod) genus. Bulletin Van Het State, Ober-devon). Göttinger Arbeiten zur Geologie und Koninkluk Belgisch Institut voor Naturwetenschappen, 73: Paläontologie, 46: 1-115, pls 1-5. 29-51. Schindler, E. 1993. Event-stratigraphic markers within the Sass, D. B. 1951. Paleoecology and Stratigraphy of the Genun- Kellwasser crisis near the Frasnian/Famennian boundary HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 201

(Upper Devonian) in Germany. Palaeogeography, Paleo- Tesmer, I. H. 1954. Geology of Chautauqua County, New climatology, Palaeoecology, 104: 115-125. York. Hobbies Magazine, Buffalo Society of Natural Sci- Schmidt, H. 1935. Die bionomische Eintelung der fossilen ences, 35: 28-35. Meeresboden. Fortschritte in der Geologie und Palaeon- Tesmer, I. H. 1975. Geology of Cattaraugus County, New tologie, 12(38): 1-154. York. Buffalo Society of Natural Sciences Bulletin, 27: 1- Sedgwick, A., & R. I. Murchison. 1839a. On the classifica- 105, 14 pls. tion of the older rocks of Devon and Cornwall. Proceed- Thayer, C. W. 1974. Marine paleontology in the Upper De- ings of the Geological Society of London, 3: 121-123. vonian of New York. Lethaia, 7: 122-155. Sedgwick, A., & R. I. Murchison. 1839b. Classification of Thompson, J. B., & C. R. Newton. 1987. Ecological rein- the older stratified rocks of Devonshire and Cornwall. terpretation of the dysaerobic Leiorhynchus fauna: Upper Philosophical Magazine, series 3, (April): 241-260. Devonian Geneseo black shale, Central New York. Sparling, D. R. 1992. On the age of the Hungry Hollow Palaios, 2: 274-281. Limestone. Journal of Paleontology, 66(2): 339. Tilton, J. L. 1929. Marine faunas of the Devonian tree hori- Sparling, D. R. 1999. Conodonts from the Prout Dolomite zons of Tygart Valley, West Virginia. American Journal of of north-central Ohio and Givetian (Upper Middle De- Science, series 5, 17: 347-351. vonian) correlation problems. Journal of Paleontology, 73 Torrey, P.D., C. E. Fralich, W. H. Young, Jr., C. Brewer, Jr., (5): 892-907. & P. M. Phillipi. 1932. The geology of New York and Steininger, J. 1849. Die Versteinerungen des Ubergangs northern Pennsylvania. American Petroleum Institute, Di- Gebirges der Eifel. Trier, 34 pp. vision of Production, Paper, 826 4A: 1-19. Steininger, J. 1853. Geognostische Beschreibung der Eifel. Trainer, D. W., Jr. 1932. The Tully Limestone of central Trier, 143 pp, 9 pls. New York. New York State Museum Bulletin, 291: 1-43. Strand, E. 1929. Zoological and palaeontological notes. Acta Trouessart, E. L. 1898. Diagnoses preliminaire d’espèce nou- Universitatis Latviensis, 20: 1-29. velles de Sarcoptidae plumicoles (Acar). Société de Ento- Stumm, C. E. 1951. Check list of fossil invertebrates de- mologie France Bulletin, 1898: 319-322. scribed from the Middle Devonian Traverse group of Tucker, M. E. 1973. Sedimentology and diagenesis of De- Michigan. University of Michigan, Museum of Paleontol- vonian pelagic limestones (Cephalopodenkalke) and as- ogy Contributions, 9(1): 1-44. sociated sediments of the Rhenohercynian Geosyncline, Sutton, R. G. 1960. Stratigraphy of the Naples Group (Late West Germany. Neues Jahrbuch für Geologie und Paläon- Devonian) in western New York. New York State Museum tologie, Abhandlungen, 142: 320-350. and Science Service Bulletin, 380: 1-56. Turner, S. 1998. "Dittodus" species of Eastman 1899 and Sutton, R. G. 1963. Correlation of Upper Devonian strata Hussakof and Bryant 1918 (Mid to Late Devonian). in south-central New York. Pp 87-101, in: Symposium Modern Geology, 21: 87-119. of Middle and Upper Devonian Stratigraphy of Pennsylva- Van Tassell, J. 1987. Upper Devonian Catskill Delta marine nia and Adjacent States, V. C. Shepps (ed.), Pennsylvania cyclic sedimentation: Brallier, Scherr and Foreknobs For- Geological Survey Bulletin, series 4, G-39: 87-101. mations of Virginia and West Virginia. Geological Society Sutton, R. G., Z. P. Bowen, & A. L. McAlester. 1970. Ma- of America Bulletin, 99: 414-426. rine shelf environments of the Upper Devonian Sonyea Vanuxem, L. 1839. Third annual report of the geological group of New York. Geological Society of America Bul- survey of the Third District. New York Geological Survey letin, 81: 2975-2992. Annual Report, 3: 241-285. Sutton, R. G., E. C. Humes, Jr., R. C. Nugent, & D. L. Vanuxem, L. 1842. Geology of New York. Part III. Comprising Woodrow. 1962. New stratigraphic nomenclature for the Survey of the Third Geological District. White and Vis- Upper Devonian of south-central New York. American sher, Albany, 307 pp. Association of Petroleum Geologists Bulletin, 46: 390-393. Weary, D. J., & A. Harris. 1994. Early Frasnian (Late De- Sutton, R. G., & G. R. McGhee, Jr. 1985. The evolution of vonian) conodonts from the Harrell Shale, Western Frasnian marine “community-types” of south-central Foreland Fold-and-thrust Belt, West Virginia, Maryland, New York. Pp 211-224, in: The Catskill Delta,D.L. and Pennsylvania Appalachians, U. S. A. Courier Woodrow & W. D. Sevon (eds), Geological Society of Forschungsinstitut Senckenberg, 168: 195-225. America Special Paper 201, Boulder, Colorado. Wedekind, R. 1913. Die Goniatitenkalke des unteren Termier, H., & G. Termier. 1950. Paléontologie Marocaine, Oberdevon von Martenberg bei Adorf. Sitzungberichte Invertebrès de l'Ere Primaire, Fascicule 3, Mollusques. Her- der Gesellschaft Naturforschender Freunde zu Berlin, 1: 1- mann, Paris, 246 pp, pls 123-183. 77, pls 1-4. 202 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

Wedekind, R. 1914. Monographie der Clymenien des ologists Memoir 14. Rheinischen Gebirges. Abhandlungen der Königlichen Work, D. M., C. E. Mason, & G. Klapper. 2007. The Mid- Gesellschaft der Wissenschaften zu Göttingen, Mathema- dle Devonian (Givetian) ammonoid Pharciceras from the tisch-physikalische Klasse, Neue Folge, 10: 1-73, pls 1-7. New Albany Shale, Kentucky. Journal of Paleontology, Wedekind, R. 1918 (1917 preprint valid for taxonomic pur- 81(6): 1510-1515. poses). Die Genera der Palaeoammonoidea (Goniatiten). Xing Li, Y. 2000. Famennian tentaculitids of China. Journal Palaeontographica, 62: 85-184, 9 pls. of Paleontology, 74: 969-975. Wells, J. W. 1956a. Receptaculites in the Upper Devonian of Xu, Y. S. 1977. Anthozoa (in Chinese). Pp 109-270, in: An- New York. Journal of Paleontology, 30(3): 747-748. thozoa. Palaeontological Atlas of Central and South China, Wells, J. W. 1956b. The ammonoids Koenenites and Beloceras H. Z. Jia, Y. S. Xu, G. D. Kuang, B. F. Zhang, Z. B. in the Upper Devonian of New York. Journal of Paleon- Zuo, & J. Z. Wu (eds), Hubei Institute of Geosciences, tology, 30: 749-751. 2, Geological Publishing House, Beijing. Wells, J. W. 1963. Early investigations of the Devonian Sys- Yatskov, S. V. 1990. K sistimatike semeistva Beloceratidae tem in New York, 1656-1836. Geological Society of Amer- (Ammonoidea). Trudy Paleontologicheskogo Instituta, ica Special Paper, 74: 1-74. 243: 36-51. Weyer, D. 1972. Zum Alter der Ammonoideen Fauna des Yochelson, E. L., & W. Kirchgasser. 1996. The youngest Marshall Sandsteins (Unterkarbon; Michigan, U.S.A.). styliolines and nowakiids (Late Devonian) currently Berichte der Deutschen Gesellschaft Geologie Wissenschaft, known from New York. Journal of Paleontology, 60(3): A, Geologie und Paläontologie, 17: 325-350. 689-700. White, I. C. 1881. The Geology of Erie and Crawford Coun- Zambito, J. J., IV, G. Baird, C. Brett, & A. J. Bartholomew. ties. Geological Survey of Pennsylvania, 2nd Report, 2007. Re-examination of the type Ithaca Formation: cor- Harrisburg, Pennsylvania, 355 pp. relations with sections in western New York. Pp 83-105, Wiedman, J., & J. Kullmann. 1981. Ammonoid sutures in in: Field Trip Guidebook for the 79th Annual Meeting of ontogeny and phylogeny. Pp 215-255, in: The Am- the New York State Geological Association, September 28- monoidea, The Evolution, Classification, Mode of Life and 30, 2007, Department of Geology, State University of New Geological Usefulness of a Major Fossil Group,M.R. York Cortland, C. A. McRoberts, (ed.), New York State House & J. R. Senior (eds), The Systematics Association Geological Association, Albany, New York. Special Volume 18, Academic Press, London. Zambito, J. J., IV, G. Baird, C. E. Brett, & A. J. Bartholo- Williams, G. Q. 1951. The Stratigraphy of the Type Ithaca mew. Unpublished data. Depositional sequences within Formation. Unpublished Ph. D. Dissertation, Cornell the type Ithaca Formation (Upper Devnian-Frasnian) in University, Ithaca, New York, 118 pp. the northern Appalachian Basin of New York State. Williams, H. S. 1884. On the fossil faunas of the Upper De- Manuscript in preparation. vonian. United States Geological Survey Bulletin, 3: 55- Ziegler, W. 1962. Taxionomie und Phylogenie oberdevonis- 86 (5-36). cher Conodonten und ihre stratigraphische Bedeutung. Williams, H. S. 1887. On the fossil faunas of the Upper De- Abhandlungen des Hessischen Landesamtes für Boden- vonian; the Genesee section, N.Y. United States Geolog- forschung, 38: 1-166, 14 pls. ical Survey Bulletin, 41: 1-123. Ziegler, W. 1971. Conodont stratigraphy of the European Williams, H. S. 1913. Recurrent Tropidoleptus zones of the Devonian. Pp 227-284, in: Symposium on Conodont Upper Devonian in New York. United States Geological Biostratigraphy, W. C. Sweet & S. M. Bergström (eds), Survey Professional Paper, 79: 1-103. Geological Society of America Memoir 127, Boulder, Wilson, L. G. 1998. Lyell in America: Transatlantic Geology, Colorado. 1841-1853. Johns Hopkins University Press, Baltimore, Ziegler, W., & C. A. Sandberg. 1990. The Late Devonian Maryland, 429 pp. standard conodont zonation. Courier Forschungsinstitut Woodrow, D. L., J. M. Dennison, F. R. Ettensohn, W. T. Senckenberg, 121: 1-115. Sevon, & W. T. Kirchgasser. 1989. Middle and Upper Ziegler, W., & C. A. Sandberg. 1994. Conodont phyloge- Devonian stratigraphy and paleogeography of the central netic-zone concept. Newsletters on Stratigraphy, 30: 105- and southern Appalachians and eastern midcontinent, 123. U. S. A. Pp 277-301, in: Devonian of the World, Proceed- Ziegler, W., & C. A. Sandberg. 1996. Reflections on Frasn- ings of the Second International Symposium on the Devon- ian and Famennian stage boundary decisions as a guide ian System, Calgary, vol. 1, N. J. McMillan, A. F. Embry, to future deliberations. Newsletters on Stratigraphy, 33: & J. D. Glass (eds), Canadian Society of Petroleum Ge- 157-180. HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 203

PLATES 204 BULLETINS OF AMERICAN PALEONTOLOGY no. 374 HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 205

PLATE 1

Figure Page 1–8. 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, 1 mi (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 near Lodi 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, West Brook 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 the West Brook Member of the Tully Limestone, 1 mi (1.6 km) northeast of Laurens, Otsego County (Loc. 9b), X 0.8.

9. Wellsites tynani House&Kirchgasser,1993...... 96 SUI 42418, holotype from the Moreland Shale (basal tongue of the Rhinestreet Shale) found in spoil from a roadcut for US Rte. 17 at Elmira, Chemung County (Loc. 70), X 0.8. Photograph by M. C. Tynan.

10. Pharciceras tridens (G. & F. Sandberger, 1849) ...... 90 Wiesbaden 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 Page 1-2. 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...... 96 NYSM 12025, lateral view of a specimen from the "Chemung" between Chemung and North Chemung, Chemung County (Loc. 95), X 0.8.

4-7. Sandbergeroceras syngonum (Clarke, 1897) ...... 93 4-5. 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) ...... 97 NYSM 4073, lateral view of the holotype figured by Vanuxem (1842: text-fig. 49, fig. 1) probably from the Cayuta Shale (Rhinestreet Shale equivalent), "near Owego," Tioga County (Loc. 69), X 1.6. HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 207 208 BULLETINS OF AMERICAN PALEONTOLOGY no. 374 HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 209

PLATE 3

Figure Page 1. Schindewolfoceras? aff. equicostatum (Hall, 1874) ...... 98 NYSM 12027, latex mold of a specimen from the "Upper Cayuta Chesney Formation" (middle Rhinestreet Shale equivalent) 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...... 95 NYSM 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) ...... 97 NYSM 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) ...... 93 5. 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; probably upper 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. 210 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

PLATE 4

Figure Page 1-4. 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 float on 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)...... 99 5. 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, Tompkins County, 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 of the 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, Tompkins County, 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. HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 211 212 BULLETINS OF AMERICAN PALEONTOLOGY no. 374 HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 213

PLATE 5

Figure Page 1-9. Ponticeras perlatum (Hall,1874)...... 99 1-4. NYSM 13082, specimen from the lower part of the Sherburne Siltstone (Lodi Limestone) in Lodi Glen, Seneca Lake (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, Seneca County, 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), near Lodi, Seneca County, J. W. Wells Coll. 400117. 7, X 0.8; 8-9, X 1.2. 214 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

PLATE 6

Figure Page 1-15. Acanthoclymenia neapolitana (Clarke, 1892) ...... 103 All specimens are baritic shell replacements apparently from the level of the Shurtleff Septarian Horizon in the Cashaqua 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. HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 215 216 BULLETINS OF AMERICAN PALEONTOLOGY no. 374 HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 217

PLATE 7

Figure Page 1. 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) ...... 160 SM XI 320c, whorl cross section and lateral views of a specimen collected by MRH from the Frasnian Büdesheimer Schiefer about 150 m southwest of the church at Büdesheim, Eifel, West Germany, X 6.

4-6. Prochorites alveolatus (Glenister, 1958) ...... 109 NYSM 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. 218 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

PLATE 8

Figure Page 1-4, 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 surviving syntype; recorded by Clarke as from the Naples Beds, Naples, Ontario County; probably from the upper Penn Yan Shale, X 1.2. 3. NYSM 12134, lateral view showing growth lines and ventrolateral furrow, Upper Penn Yan Shale, Linden Goniatite Horizon, 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), Keuka Lake, 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 Penn Yan 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...... 119 5. 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...... 121 10-11. 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. HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 219 220 BULLETINS OF AMERICAN PALEONTOLOGY no. 374 HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 221

PLATE 9

Figure Page 1. 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) ...... 130 NYSM 3804, ventral and lateral views of the lectotype, selected herein, a plastotype of the surviving syntype figured by Clarke (1898: pl. 6, fig. 31) recorded as from the Naples Beds at Naples and by Miller (1938: 123) as from the Cashaqua Shale in Briggs Gully (Loc. 43), Honeoye Lake, Ontario County, X 3.7.

4-5. Manticoceras simulator (Hall, 1874) ...... 124 NYSM 3797, ventral and lateral views of the holotype figured by Hall (1876: pl. 69, figs 1-2) and recorded as from Ithaca, Tompkins County; probably from the Ithaca Formation of the Genesee Group, X 1.2.

6-8. Manticoceras sinuosum sinuosum (Hall, 1843) ...... 124 NYSM 3762, enlarged lateral view and lateral and ventral views of a specimen figured by Clarke (1898: pl. 1, fig. 10, pl. 4, fig. 16) as Mant. Pattersoni, recorded as from "a concretion in the soft shales at Naples, N.Y.," and thought to be from the Cashaqua Shale. 6, X 1.6; 7, X 0.8; 8, X 0.6. 222 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

PLATE 10

Figure Page 1-12. Manticoceras sinuosum sinuosum (Hall, 1843) ...... 124 1-2. NYSM 13805 (CUMP 42004), apertural and lateral views of specimen collected by Donald Zenger from the Parrish 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, upper Cashaqua 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. HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 223 224 BULLETINS OF AMERICAN PALEONTOLOGY no. 374 HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 225

PLATE 11

Figure Page 1-13. 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. 226 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

PLATE 12

Figure Page 1-7, 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 baritized from the lower Cashaqua Shale, Whetstone Brook (Loc. 42/5), near Honeoye, Ontario County. 1, X 0.8; 2, X 1.6. 3-4. NYSM 13839, lateral view of the holotype with the same preservation as specimen in Figs 1-2, from the lower Cashaqua 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 Cashaqua Shale, 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) as that in Fig 6, X ca. 12.

8-10. Manticoceras sinuosum sinuosum (Hall, 1843) ...... 124 NYSM 13829, lateral, ventral, and apertural views of baritized specimen from the Shurtleff Septarian Horizon, upper Cashaqua Shale, Shurtleff's Gully (Loc. 41/1), near Livonia, Livingston County, X 1.2. HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 227 228 BULLETINS OF AMERICAN PALEONTOLOGY no. 374 HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 229

PLATE 13

Figure Page 1-15. 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. 230 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

PLATE 14

Figure Page 1-10. 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) as Manticoceras pattersoni and show- ing the protoconch; recorded as from the Naples Beds, Honeoye Lake (not Naples, New York) probably from the Cashaqua 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. HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 231 232 BULLETINS OF AMERICAN PALEONTOLOGY no. 374 HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 233

PLATE 15

Figure Page 1-2. Manticoceras sinuosum sinuosum (Hall, 1843) ...... 124 MRH Collection, baritized replacement of the shell showing dorsal wrinkle layer and its confluence with the ventral wrinkle 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. 234 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

PLATE 16

Figure Page 1-2, 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 of Farnham 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) ...... 136 All 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) ...... 124 11. 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. HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 235 236 BULLETINS OF AMERICAN PALEONTOLOGY no. 374 HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 237

PLATE 17

Figure Page 1. 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) ...... 85 2. 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...... 137 NYSM 3730, lateral view of a specimen figured by Clarke (1898: pl. 6, fig. 10) as Manticoceras accelerans and recorded as from the "soft shales at Naples," Ontario County, X 3.2.

5-10. Manticoceras sinuosum apprimatum Clarke (1898)...... 123 5. 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 as that in Figs 6-7, X 2.5. 238 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

PLATE 18

Figure Page 1-2, 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 to be 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 to be from the Genundewa Limestone on Canandaigua Lake, Yates County, X 3.8.

3-4, 6-9. Carinoceras sororium (Clarke, 1898) ...... 137 3-4. 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 the lower Angola Shale, X 3.8.

10. Delphiceras unduloconstrictum (Miller, 1938) ...... 141 GSC 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) ...... 123 USNM 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. HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 239 240 BULLETINS OF AMERICAN PALEONTOLOGY no. 374 HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 241

PLATE 19

Figure Page 1-9. 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 the Hanover Shale near Java, Wyoming County, and probably from nodules in the lower Hanover Shale in Beaver Meadow 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 as the specimen in Fig. 1, X 4. 5. NYSM 3639, lateral view of a cotype figured by Clarke (1898: pl. 6, fig. 4) from the same locality and horizon as the specimen in Fig. 1, X 4. 6. NYSM 3642, lateral view of a cotype figured by Clarke (1898: pl. 6, fig. 7) from the same locality and horizon as the specimen in Fig. 1, X 4. 7. NYSM 3640, lateral view of a cotype figured by Clarke (1898: pl. 6, fig. 5) from the same locality and horizon as the 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 locality and horizon as the specimen in Fig. 1, X 4.

10-16. Manticoceras contractum Clarke, 1898 ...... 123 10-11. 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. 242 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

PLATE 20

Figure Page 1-12. Sphaeromanticoceras rhynchostomum (Clarke, 1898) ...... 142 1-2. NYSM 12078, oblique and lateral stereoscan photographs of a baritic shell replacement of inner whorls based on a specimen from the Point Breeze Goniatite Bed, Angola Shale, in Hampton Brook (Loc. 74/6b), Erie County. 1, X8;2,X16. 3-6. NYSM 12079, oblique, ventral, and apertural views of a baritic shell replacement of inner whorls of a specimen from 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 upper Rhinestreet 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 the West 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. HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 243 244 BULLETINS OF AMERICAN PALEONTOLOGY no. 374 HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 245

PLATE 21

Figure Page 1-12. Sphaeromanticoceras rhynchostomum (Clarke, 1898) ...... 142 1-2. NYSM 3789, ventral and lateral views of a gutta percha mold figured by Clarke (1898: pl. 4, fig. 11) prepared from a specimen from Big Sister Creek (Loc. 73), Angola, Erie County, and probably from the lower Angola Shale, X 3.8. 3-4. NYSM 3786, lateral and ventral views of a gutta percha mold figured by Clarke (1898: pl. 4, fig. 8) prepared from 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 same locality 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 of Figs 1-2, X 3.7. 246 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

PLATE 22

Figure Page 1-9. Sphaeromanticoceras rhynchostomum (Clarke, 1898) ...... 142 1-3. NYSM 12089, ventral, lateral, and cross sectional views of a specimen from the lower Angola Shale in Hampton Brook (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 the Naples Beds at Angola, and probably from the lower Angola Shale at Big Sister Creek (Loc. 73), Angola, Erie County, X 0.8. 6-7. NYSM 3796, lectotype, here designated, a cotype figured by Clarke (1898: 68, text-fig. 43) probably from the lower 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), Erie County, X 0.8. 9. NYSM 12085, cross section showing the form of the early whorls and an asymmetrically placed nepionic whorl based on a specimen from upper Rhinestreet Shale equivalents in Johnson Creek (Loc. 58/1), Wyoming County, X 6.4. HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 247 248 BULLETINS OF AMERICAN PALEONTOLOGY no. 374 HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 249

PLATE 23

Figure Page 1-4. 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)...... 138 5-6. 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 Stony Creek (Loc. 83/8), Warsaw, Wyoming County, X 1.2. 250 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

PLATE 24

Figure Page 1-4. 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, Ontario County, figured by Clarke (1898: pl. 8, fig. 3), possibly also from Genundewa Point, Canandaigua Lake, as stated on 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 Genundewa Limestone at Genundewa Point (Loc. 23b), Canandaigua Lake, Yates County, X 8.

5-9. Probeloceras lutheri (Clarke, 1885)...... 156 5, 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), near Naples, 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 as from "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. HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 251 252 BULLETINS OF AMERICAN PALEONTOLOGY no. 374 HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 253

PLATE 25

Figure Page 1. 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, X 3.6.

2-5. Naplesites iynx (Clarke, 1898) ...... 161 All 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) ...... 160 NYSM 4072, holotype of Probeloceras? naplesense (Clarke, 1898), sometimes assigned to Neomanticoceras. Designated type species of Naplesites by Yatskov (1990). Figured by Clarke (1898: pl. 7, fig. 18), and probably from the Rhinestreet Shale at Naples, Yates County, X 1.2. 254 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

PLATE 26

Figure Page 1-4, 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, Livingston County, X 2.4.

5. Tornoceras uniangulare widderi House,1965...... 81 USNM 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 ...... 165 7. 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. HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 255 256 BULLETINS OF AMERICAN PALEONTOLOGY no. 374 HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 257

PLATE 27

Figure Page 1-6. 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 Windom Shale at Grove's Creek Quarry (Loc. 4), southeast of Ovid, Seneca County, X 3.2.

7-10. Truyolsoceras bicostatum (Hall, 1843) ...... 179 NYSM 11957 (MRH D1364), lateral and ventral views of the early whorls showing the nepionic constriction and commencement of ventrolateral furrows of a specimen collected by MRH from the Corell's Point Goniatite Bed at Corell's Point (Loc. 97), near Brocton, Chautauqua County, X 16.

11. Tornoceras? sp.House,1965...... 11 NYSM 11633 (5733), lateral view of a specimen from the West Brook Member of the Tully Limestone from near Borodino (Loc. 6), Onondaga County, X 0.8.

12. Linguatornoceras aff. linguum (G. & F. Sandberger, 1851) ...... 167 NYSM 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)...... 167 13. 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) ...... 166 NYSM 4093, lateral view of a specimen from the “Naples formation” at Mount Morris, Livingston County, figured by Hall (1879: pl. 72, figs 6-7), X 1.2.

16. Tornoceras arcuatum House, 1965 ...... 165 NYSM 12671 (MRH D1440), lateral view of a topotype collected by MRH from the Squaw Bay Limestone near Partridge Point, Alpena, Michigan, X 2.4. 258 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

PLATE 28

Figure Page 1-5. 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 Bethany Center (Loc. 24), Genesee County, X 1.2.

6-10. Tornoceras uniangulare obesum Clarke, 1897 ...... 165 6-7, 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, Ontario County, X 3.9. HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 259 260 BULLETINS OF AMERICAN PALEONTOLOGY no. 374 HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 261

PLATE 29

Figure Page 1-15. Phoenixites concentricus (House, 1965)...... 173 1, 4-5. NYSM 11964 (6679E), ventral, lateral, and apertural views of a specimen from the Corell's Point Goniatite Bed 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 and development thereafter of ventrolateral furrows, based on a specimen from the Corell's Point Goniatite Bed at Corell'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, Gowanda Shale, 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 as that 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 at Corell'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 locality and horizon as that in Fig. 13, X 2.4. 262 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

PLATE 30

Figure Page 1, 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 or Westfield 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 or Westfield Shale, X 1.2.

2, 4-14. Truyolsoceras bicostatum (Hall, 1843) ...... 179 2. 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 at Corell'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 Goniatite Bed (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)...... 180 NYSM 12120, lateral view of a specimen from the Gowanda Shale from near the Mixer Road bridge over Walnut Creek (Loc. 99a) at Forestville, Chautauqua County, X 1.6. HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 263 264 BULLETINS OF AMERICAN PALEONTOLOGY no. 374 HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 265

PLATE 31

Figure Page 1. Truyolsoceras cf. bicostatum (Hall, 1843)...... 180 NYSM 12119, lateral view of a specimen from the Gowanda Shale from near the Mixer Road bridge over Walnut Creek (Loc. 99a) near Forestville, Chautauqua County, X 1.6.

2-3. Aulatornoceras auris (Quenstedt, 1846) Group ...... 176 2. 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) ...... 177 4. 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) ...... 176 7. 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...... 92 14. 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. 266 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

PLATE 32

Figure Page 1. Aulatornoceras rhysum (Clarke, 1898) ...... 179 NYSM 4092, lateral view of the lectotype figured by Clarke (1898: text-fig. 100, pl. 8, fig. 14) labeled “Hanover Shale, Java” and probably from the lower Hanover Shale in Beaver Meadow Creek at Java (Loc. 92), Wyoming County, X 4.

2-13. Aulatornoceras paucistriatum (d'Archiac & de Verneuil, 1842)...... 178 2-3. 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) ...... 177 NYSM 12102, lateral view of a specimen from near the Point Breeze Goniatite Bed of the Angola Shale in Hampton Brook (Loc. 74/6a), X 3.2. HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 267 268 BULLETINS OF AMERICAN PALEONTOLOGY no. 374 HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 269

PLATE 33

Figure Page 1-2, 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 Tonowanda Creek 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 Johnson Creek (Loc. 58/1), Wyoming County, X 2.

3-4. Linguatornoceras sp.,juvenilespecimen...... 167 NYSM 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)...... 168 9-10. 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. 270 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

PLATE 34

Figure Page 1-4. 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) ...... 171 CU 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) ...... 171 NYSM/CU 369, lateral and ventral views of a specimen from the upper Geneseo Shale or lower Penn Yan Shale, Fir Tree Point, western side of Seneca Lake (Loc. 25), Yates County, X 0.8.

8. Epitornoceras peracutum (Hall, 1876) ...... 170 NYSM 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. HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 271 272 BULLETINS OF AMERICAN PALEONTOLOGY no. 374 HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 273

PLATE 35

Figure Page 1-10. 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 Erie Shore, 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)...... 183 USNM 137645, lateral view of a specimen collected by H. S. Williams in 1884 from the Ellicott Shale at Porter Creek, Summerdale (Loc. 110), Chautauqua County. 11, X 0.4; 12, X 0.7.

12. ?Genus edwinhalli (Clarke, 1898) ...... 183 NYSM 4090, lateral view of the holotype figured by Clarke (1898: 111, text-fig. 85) from the Conneaut Formation at Nile, Allegany County, X 1.6. 274 BULLETINS OF AMERICAN PALEONTOLOGY no. 374 HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 275

SYSTEMATIC INDEX Pages denoting the start of principal discussions, locations of plate and text-figure illustrations, and tables of measurements are in bold font.

Acanthoclymeniidae Schindewolf, 1955...... arcuatum, Tornoceras ...... 22,79, 82, 158, 165-166, 17-, 254, 257; ...... 72-73, 77, 98, 103, 114, 155; Text-figs 20-21 Text-figs 23, 54H-J, L; Pl. 26, Figs 7-10, Pl. 27, Fig. 16 Acanthoclymenia Hyatt, 1900...... 17,21,23,29-30,32, arcuatum, Tornoceras cf...... 10,158,166,170;Text-fig. 54K 72-73, 82, 98, 102-104, 107, 109, 120, 157; Text-figs 21, 30 arkonense, Tornoceras uniangulare (House, 1965) ...... 81 A. forcipifer (G.&F.Sandberger,1850)...... 102, 247 astarte, Goniatites ...... 163 A. genundewa (Clarke,1898)...... 79, 105-107, 155, Atlantoceras Bensaïd, 1974 ...... 76, 93, 102 250; Text-figs 23, 30A-C, K, L; Table 6; Pl. 24, Figs 1-4 Aulatornoceratini Becker, 1993...... 175 A. aff. genundewa (Clark, 1898) ...... 106 Aulatornoceras Schindewolf, 1922 ...... A. cf. genundewa (Clarke, 1898) ...... 105-106, 107 ...... 39-40, 73, 86, 168, 175, 178-179; Text-fig. 21 A. neapolitana Clarke, 1892)...... 35,39-40, 79, 83, 103- A. eifliense (Steininger, 1849) ...... 106, 214; Text-figs 23, 30E-H; Table 5; Pl. 6, Figs 1-15 ...... 35,38-40, 79, 83, 175-176, 177-178, A. aff. neapolitana (Clarke, 1892) ...... 265; Text-fig. 23; Table 31; Pl. 31, Figs 7-13, 16-20 ...... 31,33,79, 83, 102-105; Text-figs 23, 30D A. aff. eifliense (Steininger, 1849)....49,54,79, 177, 265-266; A. cf. neapolitana (Clarke, 1892) ...... 106 Text-fig. 23; Table 32; Pl. 31, Figs 4-6, Pl. 32, Fig. 14 A. orientale (Bogoslovsky, 1969) ...... 102 A. eifliense eifliense (Steininger, 1849)...... 177 A. planorbe (G. & F. Sandberger, 1850)...... 102 A. eifliense posterior Becker, 1993 ...... 175 A. sp...... 79, 103, 107; Text-fig. 23 A. auriforme (Oppenheimer, 1916) ...... 176 A. sp. A Kirchgasser, 1975 ...... 31-32, 105 A. auris (Quenstedt, 1846)...... 55,79, 85, accelerans, Manticoceras ...... 1, 137, 237 175-176, 178, 265; Text-figs 23, 60G; Pl. 31, Figs 2-3 acutiforme, Carinoceras ...... 137-138, 140 A. cf. auris (Quenstedt, 1846)...... 176 acutolaterale, Maeneceras ...... 86,183, 185 A. auris bickensis (Wedekind, 1918) ...... 175 acutolaterale, Maeneceras aff...... 62,79, 86, 182-183, 184- A. constrictum (Steininger, 1853)...... 176 185, 273; Text-figs 23, 61C-D, 62-63; Pl. 35, Figs 11, 13 A. keyserlingi (Müller, 1956) ...... 176 acutolateralis, Goniatites ...... 182 A. lepiferum Becker, 1993 ...... 175 acutum, Carinoceras ...... 137-138, 140; Text-fig. 44B A. loeschmanni (Frech, 1902) ...... 169, 176 acutum, Oxytornoceras ...... 169 A. paucistriatum (d’Archiac & de Verneuil, 1841) ...... acutum, Tornoceras ...... 169 ...... 54-55,79, 85, 175-176, 178-179, 180, aequabile, Ponticeras ...... 98 266; Text-figs 23, 60D, F; Table 33; Pl. 32, Figs 2-13 affine, Sphaeromanticoceras ...... 142, 157 A. rhysum (Clarke, 1898) ...... 58-59, 79, 85, affinis, Goniatites ...... 142 175-176, 178-179, 266; Text-figs 23, 60E; Pl. 32, Fig. 1 Afromaenioceras Göddertz,1987...... 73, 76; Text-fig. 21 A. sandbergeri (Ford & Crick, 1897)...... 175 altaicum, Manticoceras ...... 150 A. serriense Becker, 1993 ...... 176 alveolatus, Prochorites ...... 35, 38-41, 79, 83, 109-110, 156-157, auriforme, Aulatornoceras ...... 176 185, 217; Text-figs 23, 31, 32C, 63; Table 7; Pl. 7, Figs 4-6 auris, Aulatornoceras ...... 55,79, alveolatus, Prochorites aff...... 43,49,110 85, 175-176, 178, 265; Text-figs 23, 60G; Pl. 31, Figs 2-3 amblylobum, Cheiloceras (Cheiloceras)...... 59-62,79,86,181-182, 185, auris, Aulatornoceras cf...... 176 273; Text-figs 23, 59C, 61A-B; Table 35; Pl. 35, Figs 1-10 auris, Goniatites ...... 175-176 Ammonites aequabile Beyrich, 1837 ...... 98 auritum, Ponticeras ...... 98 Ammonites hoeninghausia vonBuch,1832...... 112, 122 ausavense, Lobotornoceras ...... 168, 171-172; Text-figs 58C, F Ammonites orbiculus Beyrich, 1837 ...... 142 ausavensis, Goniatites ...... 168 Ammonites primordalis vonBuch,1832...... 142 barnetti, Pharciceras ...... 186 Ammonites simplex vonBuch,1832...... 163 barroisi, Ponticeras ...... 98 amplexum, Pharciceras ...... 10-11, 79, 82, 89-90, 91, 185- beckeri n. sp., Koenenites . . . . 1, 31-34, 79, 112-115, 117, 121-122, 185, 186, 205; Text-figs 23, 26D-H, 63; Table 2; Pl. 1, Figs 1-8 218; Text-figs 23, 33A-E, 34H, K, 63; Pl. 8, Figs 10-11, 14 Anarcestidae ...... 71-72, 73, 76, 98, 162; Text-figs 20-21 beckeri,n.sp.,Koenenites,formD...... anguisellatum, Naplesites ...... 160-161 ...... 31, 33, 117, 122; Text-figs 33E, 34K annissi, Crassatornoceras ...... 168 belgicum, Tornoceras ...... 168 apprimatum, Manticoceras ...... 83,123, 132 Beloceratidae Hyatt, 1884 ...... apprimatum, Manticoceras sinuosum ...... 1,29,79,83,123, 132, ...72-73, 76-77, 94, 96, 98, 102-103, 155; Text-figs 20-21 135, 185, 237; Text-figs 23, 35B, 42F, 63; Pl. 17, Figs 5-10 Beloceras Hyatt,1884...... 50,73, 83-84, 94, 96, 155; Text-fig. 21 archiaci, Hoeninghausia ...... 122 B. sagittarium (G. & F. Sandberger, 1851) . . . . 94; Text-fig. 27C archiaci, Hoeninghausia aff...... 83 B. williamsi Wells, 1956...... 96 archiaci, Hoeninghausia cf...... 121 bicaniculatum, Lobotornoceras ...... 173 Archoceras Schindewolf, 1937...... 79, 83, 85, 102, 149; Text-fig. 48 bickensis, Aulatornoceras auris ...... 175 A. varicosum (Drevermann, 1901) ...... 85 bicostatum, Tornoceras ...... 178 A. wabashense (Kindle, 1901) ...... 237; Pl. 17, Fig. 1 bicostatum, Tornoceras (Aulatornoceras)...... 178 A. (Atlantoceras) sp. Kirchgasser & House, 1981...... 102 bicostatum, Truyolsoceras ...... ?A. sp...... 58-59, 79, 86, 155, 192; Text-figs 23, 48G, 63 . . . . . 1, 59-61, 79, 86, 175-176, 178-179, 180, 257, 262; Text- Atlantoceras Bensaïd, 1974 ...... 76,98,102 figs 23, 59B, 60A-C, H; Table 34; Pl. 30, Figs 2, 4-13, 15 276 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

bicostatum, Truyolsoceras cf...... 59-61, 79, concentricum, Tornoceras (Tornoceras)...... 173 86, 180, 262, 265; Text-fig. 23; Pl. 30, Fig. 15, Pl. 31, Fig. 1 concentricus, Phoenixites ...... 59-60, 86, 158, bidentatum, Pharciceras ...... 89 173, 261; Text-figs 54M-O, 59A; Table 26; Pl. 29, Figs 1-15 bilobatum, Falcitornoceras aff...... 172; Text-fig. 58D constrictum, Aulatornoceras ...... 176, 179 bisulcatum, Ponticeras ...... 98 contractum, Manticoceras ...... 1,27,29,79,83,118,123, buchii, Manticoceras ...... 129 130, 132, 136-137, 241; Text-figs 23, 35B; Pl. 19, Figs 10-16 Carinoceras Lyashenko,1957...... cooperi, Koenenites ...... 112-113, 117, 120; Table 9 ...... 73, 77, 84, 122, 129, 137, 140; Text-figs 21, 42-44 cordatum, Manticoceras ...... 129, 134, 136 C. acutiforme (H.&G.Termier,1950)...... 137-138,140 cordatum, Manticoceras lamed aff...... C. acutum (G. & F. Sandberger, 1850)...... 31,83,115, 136; Text-figs 33K-L ...... 137-138, 140; Text-fig. 44B cordatum, Manticoceras lamed cf...... 83 C. galeatum (Wedekind, 1913)...... 85,140 cordiforme, Crickites ...... 155 C. menneri Ljaschenko, 1957 ...... 137-138 cordiforme, Manticoceras ...... 129, 150 C. oxy (Clarke, 1897)...... 144 Costamanticoceras Becker & House, 1993 . . . . . 73, 77, 122; Text-fig. 21 C. sororium (Clarke, 1898) ...... 1,51,79, C. koeneni (Holzapfel, 1882)...... 137 85, 137, 238; Text-figs 23, 42G; Pl. 18, Figs 3-4, 6-9 costatum, Triainoceras ...... 94;Text-fig. 27E C. vagans (Clarke, 1898)...... costatus, Goniatites ...... 92 . . . . . 43, 48, 54-55, 79, 84-85, 138-139, 140, 249; Text- Crassotornoceras House & Price, 1985 . . . 73, 157, 162, 168; Text-fig. 21 figs 23, 43A-E, 44D, 46A; Table 16; Pl. 23, Figs 5-16 C. annissi House & Price, 1985 ...... 168 C. aff. vagans (Clarke, 1898)...... C. ausavense crassum (Matern, 1931) ...... 168 ...55,79, 138, 141, 249; Text-fig. 23; Pl. 23, Figs 1-4 C. belgicum (Matern,1931)...... 168 cataphractum, Delphiceras ...... 1,58-59,79,85,134, C. crassum (Matern, 1931) ...... 169 141, 185, 241; Text-figs 23, 41G, 63; Pl. 19, Figs 1-9 C. aff. crassum (Matern, 1931) ...... 54-57, 85, 157, cataphractum, Gephyroceras...... 141 168, 269; Text-figs 53E-F; Table 27; Pl. 33, Figs 9-20 Ceratobeloceras House & Kirchgasser, 1985 ...... 73,77,155 ?C. isolatum Becker, 1993...... 168 Chaoceras Yatskov,1991...... 161 C. nitidum Becker, 1993 ...... 168 Cheiloceras Frech, 1897 ...... 3, 59, 62, 73, 174, 181-182; Text-fig. 21 crassum, Crassotornoceras ...... 169 C. (Cheiloceras) amblylobum (G. & F. Sandberger, 1851)...... crassum, Crassotornoceras aff...... 54-57, 79, 85, 157, ...... 59-62, 79, 174, 181-182, 185; Text- 168, 269; Text-figs 23, 53E-F; Table 27; Pl. 33, Figs 9-20 figs 23, 59C, 61A-B, 63; Table 35; Pl. 35, Figs 1-10 crassum, Crassoctornoceras ausavense ...... 168 Cheiloceratidae Frech, 1897 ...... 72-73, 181; Text-figs 20-21 crassum, Tornoceras ausavense ...... 168 Cheiloceratos Strand,1929...... 181 Crickitinae ...... 122, 150 Cheilocerotes Strand,1929...... 181 Crickites Wedekind, 1913 . . . . . 73, 77, 122, 149-150, 155; Text-fig. 21 Chemungense, Sandbergeroceras ...... 97 C. cordiforme (Miller, 1938) ...... 155 chemungense, Schindewolfoceras ...... 50, C. holzapfeli Wedekind, 1913 . . . . 150, 155; Text-figs 48D-F 79, 84, 94, 96-97, 98, 206; Text-figs 23, 27I, 63; Pl. 2, Fig. 8 C. lindneri (Glenister, 1958) ...... 58, chemungense, Schindewolfoceras aff...... 79, 85, 149-150, 151, 185; Text-figs 49, 63; Table 21 ...... 94,97, 210; Text-fig. 27J; Pl. 4, Figs 1-4 ?C. sp.juv...... 79, 86, 149, 155, 185; Text-figs 23, 48H, 63 chemungensis, Goniatites ...... 93 Delphiceras Becker & House, 2000 . . 73, 77, 122, 134, 141; Text-fig. 21 Chiloceras Dreverman,1901...... 181 D. cataphractum (Clarke, 1898) ...... 1,58-59, 79, 85, 134, Chutoceras Becker&House2000...... 141, 185, 241; Text-figs 23, 41G, 63; Pl. 19, Figs 1-9 ...... 73, 98, 100, 107-109, 129; Text-fig. 21 D. unduloconstrictum (Miller, 1938) . . 141, 238; Pl. 18, Fig. 10 C. manticoides Becker & House, 2000 ...... 107-109 delphiense, Clauseniceras ...... 85,237; Pl. 17, Figs 2-3 C. nundaium (Hall, 1874)...... Delphinites Becker & House 1993 ...... 122, 141 ...... 1, 17, 23, 25-26, 77, 79, 82, 100, 107-108, Delphinites Sayn, 1901 ...... 141 109, 185, 221; Text-figs 23, 28H-K, 63; Pl. 9, Fig. 1 Devonopronorites ...... 71,72-73, 77, 89 cinctum, Goniatites ...... 176 DevonopronoritidaeBogoslovsky,1958....72-73, 89; Text-figs 20-21 clarkei, Tornoceras (Aulatornoceras) ...... 61 discoidale, Ponticeras ...... 98 clarkei, Truyolsoceras ...... 59, domanicense, Ponticeras ...... 98 61, 79, 86, 176, 179-180,262;Text-fig. 23; Pl. 30, Figs 1, 3 Domanikoceras Becker&House,1993...... 73, 162; Text-fig. 21 Clauseniceras Becker & House, 1993 ...... 73, 122; Text-fig. 21 Edwin-halli, Tornoceras ...... 62,87,183 C. delphiense (Kindle, 1901) ...... 85, 237; Pl. 17, Figs 2-3 edwinhalli,?Genus...... 183, 273; Pl. 35, Fig. 12 C. expectatum (Wedekind, 1913)...... 85 Eidoprobeloceras Kirchgasser, 1968 ...... 98 clausium n. ssp., Manticoceras sinuosum ...... E. strix Kirchgasser, 1968 ...... 109 .....1,35,40-41, 83, 129-130, 131, 226, 229; Text-figs 39A- eifliense, Aulatornoceras ...... 35,38-40, 79, 83, I, 40A-B; Table 12; Pl. 12, Figs 1-7, 11, Pl. 13, Figs 1-15 175-176, 177-178, 265; Table 31; Pl. 31, Figs 7-13, 16-20 clausum, Linguaornoceras ...... 166 eifliense, Aulatornoceras aff...... 49,54, clausum, Tornoceras ...... 166 79, 177, 265-266; Table 32; Pl. 31, Figs 4-6, Pl. 32, Fig. 14 Clymenia Münster in Goldfuss, 1832 ...... 102-103, 107-108 eifliense, Tornoceras ...... 177 complanatus perlatus, Goniatites ...... 99,108 eifliense eifliense, Aulatornoceras ...... 177 compressum, Tornoceras uniangulare ...... eifliense posterior, Aulatornoceras ...... 175 ...... 27,29,79, 82, 157-158, 162, 164-165, 258; enfieldense n. sp. , Sandbergeroceras? ...... Text-figs 23, 53A-B, 54F, 55B; Table 24; Pl. 28, Figs 1-5 . . . 1, 49-51, 93, 95, 206, 209; Pl.2,Figs1-2,Pl.3,Figs2-3 HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 277

Enseites Becker & House, 1993 ...... 73, 77, 98; Text-fig. 21 Goniatites lamellosus G. & F. Sandberger, 1850 ...... 112 E. sulcatum (Matern, 1931) ...... 105 Goniatites lingua G. & F. Sandberger, 1851 ...... 116 Eobeloceratidae ...... 71,72-73, 76, 88-89, 155; Text-figs 20-21 Goniatites lutheri Clarke, 1885 ...... 156 Epitornoceras Frech,1902...... 16,19-20, Goniatites mithracoides Frech,1887...... 169 73, 76-77, 82, 158, 162, 164, 166, 169-170, 171; Text-fig. 21 Goniatites nundaia Hall, 1874 ...... 107 E. mithracoides (Frech, 1887)....82,169-170; Text-fig. 57G Goniatites pattersoni Hall,1860...... 108 E. cf. mithracoides (Frech, 1887)...... Goniatites retrorsus var. lingua G. & F. Sandberger, 1851 ...... 166 ...... 16,24,82,170, 185; Text-figs 57F, 63 Goniatites retrorsus var. typus G.&F.Sandberger,1851...... 163 E. peracutum (Hall, 1876) ...... 74, 79, 169-170, 171, Goniatites simulator Hall,1874...... 122 185; Text-figs 23, 57A-E; Table 28; Pl. 34, Figs 5, 8 Goniatites sinuosus Hall, 1843 ...... 107-108, 124 E. aff. peracutum (Hall, 1876)...... Goniatites subpartitus Münster,1839...... 181 ...... 16,169-171; Text-figs 57B-C Goniatites (Tornoceras) subundulatus var. falcata Frech,1887...... 173 E. cf. peracutum (Hall, 1876)...... Goniatites tuberculosocostatus G. & F. Sandberger, 1850 ...... 93 ...... 16, 170-171, 185; Text-figs 57D-E, 63 Goniatites undulatus Brown,1841...... 175 equicostatum, Schindewolfoceras? ...... Goniatites undulatus G. & F. Sandberger, 1851 ...... 179 ...... 50,79, 84, 96-97, 98, 209; Text-fig. 23; Pl. 3, Fig. 4 Goniatites uniangulare Conrad,1842...... 162-163, 164 equicostatum, Schindewolfoceras? aff...... guestphalicum, Tornoceras ...... 166 ...... 50,79, 84, 96-98, 209; Text-fig. 23; Pl. 3, Fig. 1 Gundolficeras Becker,1995...... 73, 162, 173 evolutum, Manticoceras cf...... 102 hassoni, Lobotornoceras ...... 171-172; Text-fig. 58E Exotornoceras Becker,1993...... 171 hassoni, Lobotornoceras aff...... 31-32, 79, 83, 171- expectatum, Clauseniceras ...... 85 172, 270; Text-figs 23, 58A-B; Table 30; Pl. 34, Figs 1-4 falcata, Goniatites (Tornoceras) subundulatus var...... 173 haugi, Tornoceras ...... 166 Falcitornoceratini Becker, 1993 ...... 172 Hoeninghausia Gürich,1896...... 73, 83, 112, 120-122; Text-fig. 21 Falcitornoceras House & Price, 1985 ...... 73, 77, 171, 173 H. archiaci Gürich,1896...... 122 F. aff. bilobatum (Wedekind, 1908) ...... 172; Text-fig. 58D H. aff. archiaci Gürich,1896...... 83 fasciculatum, Manticoceras ...... 120 H. cf. archiaci Gürich, 1896, House et al.,1985...... 121 fasciculatus, Koenenites? ...... 1,27,29-30, 82, 112- H. (Koenenites) hoeninghausia vonBuch,1832...... 122 113, 115, 120-121, 238; Text-figs33I-J;Pl.18,Figs1-2,5 hoeninghausia, Ammonites ...... 122 fasciculatus, Koenenites? cf...... 31-32, 113, 121 hoeninghausia, Hoeninghausia (Koenenites)...... 122 Foordites Wedekind,1918...... 76,163 hoeninghausia, Koenenites ...... 112-113 forcipifer, Acanthoclymenia (G. & F. Sandberger, 1850)...... 102 holzapfeli, Crickites ...... 77, 86, 149-150, 155; Text-figs 48D-F frechi, Tornoceras ...... 173 holzapfeli, Gephyroceras ...... 105, 153, 253 galeatum, Carinoceras ...... 85,137-138, 140 housei, Naplesites ...... 160-161 galeatum, Pharciceras ...... 89,170 hunanense, Manticoceras ...... 142 Gattendorfia Schindewolf, 1920 ...... 73,92 incertum, Trianoceras ...... 92 genundewa, Acanthoclymenia ...... 1, 27-29, 82, 104-105, 106-107, inflexum, Maeneceras ...... 87,182-183; Text-fig. 61F 155-250; Text-figs 30A-C, K, L; Table 6; Pl. 24, Figs 1-4 intumescens, Manticoceras ...... 129 genundewa, Acanthoclymenia aff...... 106 inversum, Manticoceras ...... 137 genundewa, Acanthoclymenia cf...... 30,83,106-107 isolatum,?Crassotornoceras ...... 168 genundewa, Probeloceras cf...... 106 iynx, Naplesites ...... 79,83,156, 160- ?Genus edwinhalli (Clarke, 1898) ...... 183, 273; Pl. 35, Fig. 12 161, 185-186, 253; Text-figs 23, 52A-E, 63; Pl. 25, Figs 2-5 Gephuroceras Hyatt,1884...... 123 iynx, Naplesites cf...... 160-161 Gephuroceratidae Frech, 1897...... kayseri, Ponticeras ...... 98 ...... 72-73, 76-77, 98, 103, 114, 137; Text-figs 20-21 keyserlingi, Aulatornoceras ...... 176 Gephuroceratinae Frech, 1879...... 122 keyserlingi, Ponticeras ...... 98 Gephyroceras Carus, 1884 ...... 123 kilfoylei n. ssp., Koenenites styliophilus ...... 1, G. cataphractum Clarke, 1898 ...... 141, 153 27-29, 79, 82, 112, 115, 117, 119-120, 185, 218; Text-figs 23, G. holzapfeli Clarke,1898...... 105, 253 33F, G, N-O, 34G, 63; Table 10; Pl. 8, Figs 5, 7-8, 12-13 gerassimovi, Triainoceras ...... 92,94;Text-fig. 27D kirchgasseri, Koenenites lamellosus ...... 112-113, 117 Gogoceras Becker et al.,1993...... 73, 77, 98; Text-fig. 23 Kirsoceras Bogoslovsky,1971...... 73, 162 Goniatites acutolateralis G. & F. Sandberger, 1850 ...... 182 koeneni, Costamanticoceras ...... 137 Goniatites affinis Steininger,1849...... 142 Koenenitidae Becker & House, 1993 ...... Goniatites astarte Clarke,1885...... 163 ...... 72-73, 77, 98, 112, 114; Text-figs 20-21 Goniatites auris Quenstedt, 1846 ...... 175-176 Koenenites Wedekind, 1913 ...... Goniatites ausavensis Steiniger,1853...... 168 ...... 3, 13, 17-19, 21, 23, 25, 28-30, 32, 65-66, 73, 77, 82, Goniatites chemungense Clarke, 1885 ...... 93 102, 107, 112-113, 115, 117, 120-123, 129, 166; Text-fig. 21 Goniatites cinctum Münster, 1843 ...... 176 K. beckeri n.sp...... Goniatites complanatus perlatus Hall,1874...... 99,108 ...... 1,31-34, 79, 83, 112-114, 117, 121-122, 185218; Goniatites complanatus perlatus Hall,1876...... 108 Text-figs 23, 33A-E, 34H, K, 63; Pl. 8, Figs 10-11, 14 Goniatites costatus d’Archiac & de Verneuil, 1842 ...... 92 K. beckeri n. sp., form D ...... 115, 122; Text-figs 33E, 34K Goniatites lamed var. complanatus G. & F. Sandberger, 1850 . . . 132, 136 K. cooperi Miller, 1938 ...... 112-113, 117, 120; Table 9 Goniatites lamed var. latidorsalis G. & F. Sandberger, 1850 ...... 142 K.? fasciculatus (Clarke, 1898) . . 1, 27, 29, 79, 82, 112-113, 115, Goniatites lamed var. tripartitus G.&F.Sandberger,1850...... 141 120-121, 238; Text-figs 23, 33 I-J; Pl. 18, Figs 1-2, 5 278 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

K.? cf. fasciculatus (Clarke, 1898) ...... 31-32, 82, 113, 121 L. aff. hassoni ...... 31-32, 79, 83, 172, K. hoeninghausia (von Buch, 1832)...... 112-113, 122 270; Text-figs 23, 58A-B; Table 30; Pl. 34, Figs 1-4 K. lamellosus (G. & F. Sandberger, 1850)...... loeschmanni, Aulatornoceras ...... 169, 176 ...... 112-113, 117, 120-121, 122 lutheri, Probeloceras aff...... 157 K. lamellosus kirchgasseri House,1978...... 112-113, 117 lutheri, Probelcoeras cf...... 157, 160 K. aff. lamellosus (G. & F. Sandberger, 1850) sp. D House & lutheri, Goniatites ...... 156 Kirchgaser,1993...... 83,121-122 lutheri, Probeloceras ...... 35, 38-43, 79, 95, 105, K. cf. lamellosus (G. & F. Sandberger, 1850) . . . . 113, 121-122 125, 152, 155-156, 157, 159-161, 185, 250, 253; Text-figs 23, K. styliophilus (Clarke, 1898)...... 22,26,112 35C, 50-51, 63; Table 22; Pl. 24, Figs 5-9, Pl. 25, Fig. 1 K. aff. styliophilus House & Kirchgasser, 1993 ...... 119 Maeneceras Hyatt,1844...... 62,73, 87, 182-183; Text-figs 21, 61 K. styliophilus kilfoylei n.ssp...... 1,27-29, 79, M. acutolaterale (G. & F. Sandberger, 1850) ...... 86, 183, 185 82, 112, 115, 117, 119-120, 185, 218; Text-figs 23, 33F, M. aff. acutolaterale (G. & F. Sandberger, 1850) ...... G, N-O, 34G, 63; Table 10; Pl. 8, Figs 5, 7-8, 12-13 ...... 62,79, 86, 182-183, 184-185, K. styliophilus styliophilus (Clarke, 1898) . . . . 1, 17, 19-20, 22- 273; Text-figs 23, 61C-D, 62-63; Pl. 35, Figs 11, 13 23, 79, 82, 112-113, 115-117, 119-120, 185, 218; Text- M. inflexum (Wedekind, 1908) . . . . 87, 182-183; Text-fig. 61F figs 23, 34A-F, I- J, L, 35A, 63; Table 8; Pl. 8, Figs 1-9 M. milleri (Flower & Caster, 1935)...... K. styliophilus styliophilus (Clarke, 1898) morphotype B ...... 3,62,185; Text-figs 61E, 63 ...... 115, 117, 120 M. pompeckji (Wedekind, 1918) ...... 86 K. sp. House, 1978 ...... 113, 117 M. cf. pompeckji (Wedekind, 1918) ...... 62,183 K. sp. C Kirchgasser & House, 1981 ...... 121 Maenioceras Schindewolf,1933...... 76,81,90 K. sp.DKirchgasser&House,1981...... 122 MaenioceratidaeBogoslovsky,1958....71-72, 73, 76; Text-figs 20-21 Komioceras Bogoslovsky,1958...... 73, 112; Text-fig. 21 Manticoceratidae Wedekind, 1913 ...... 122 kwangsiense, Manticoceras ...... 142 Manticoceras Hyatt,1884...... kweipingense, Manticoceras ...... 142 ...... 2,43,51,73, 76-77, 83, 102-103, 105, 107, 112-113, lamed, Manticoceras ...... 115, 120, 122-124, 129, 135, 137, 142, 150, 157; Text-fig. 21 ...... 27,43,48-49, 54-55, 57, 79, 85, 132, 134-135, 234; M. accelerans Clarke, 1887...... 1,137, 237 Text-figs 23, 41A-C, 42E; Table 14; Pl. 16, Figs 1-2, 9-10 M. altaicum Bogoslovsky, 1958 ...... 150 lamed, Manticoceras aff...... 54, 57, 79, 85, 134- M. apprimatum Clarke, 1898 ...... 123, 132 136, 234; Text-figs 23, 41D-E; Table 15; Pl. 16, Figs 3-8 M. buchii (d’Archiac & de Verneuil, 1842) ...... 129 lamed aff. cordatum, Manticoceras ...... M. contractum Clarke, 1898 . . 1, 27, 29, 79, 83, 118, 123, 130, ...... 31,79, 83, 115, 136; Text-figs 23, 33K-L 132, 136-137, 241; Text-figs 23, 35B; Pl. 19, Figs 10-16 lamed cf. cordatum, Manticoceras ...... 83 M. cordatum (G. & F. Sandberger, 1850) . . 129, 134, 136 lamellosus, Koenenites aff...... 83,121-122 M. cordiforme Miller, 1938 ...... 129, 150 lamellosus, Koenenites aff.,sp.D...... 122 M. evolutum Petter, 1959 ...... 102 lamellosus, Koenenites cf...... 113, 121-122 M. cf. evolutum House&Kirchgasser,1993...... 102 lamellosus, Goniatites ...... 112 M. fasiculatum Clarke,1898...... 120 lamellosus, Koenenites ...... 112-113, 117, 120-121, 122 M. hunanense Xu,1977...... 142 latidorsalis, Goniatites lamed var...... 142 M. intumescens (Beyrich, 1837)...... 129 lepiferum, Aulatornoceras ...... 175 M. inversum Wedekind,1913...... 137 lindneri, Crickites ...... M. kwangsiense Chao, 1956 ...... 142 ....58,79, 85, 149-150, 151, 185; Text-figs 23, 49; Table 21 M. kweipingense Chao, 1956 ...... 142 lindneri, Manticoceras ...... 150 M. lamed (G. & F. Sandberger, 1850) ...... lingua, Goniatites ...... 166 27, 43, 48-49, 54-55, 57, 79, 85, 132, 134-135, 234; Text- lingua, Goniatites retrorsus var...... 166 figs 23, 41A-C, 42E; Table 14; Pl. 16, Figs 1-2, 9-10 Linguatornoceras House, 1965 ...... 73, 162, 166, 168; Text-fig. 21 M. lamed aff. cordatum (G. & F. Sandberger, 1850) ...... L. clausum (Glenister, 1958)...... 167 ...... 31,79, 83, 115, 136; Text-figs 23, 33K-L L. linguum (G. & F. Sandberger, 1851) ...... M. aff. lamed (G. & F. Sandberger, 1850) ...... 167, 257; Pl. 27, Figs 13-14 ...... 54,57,79, 85, 134-136, L. aff. linguum (G. & F. Sandberger, 1851)...... 234; Text-figs 23, 41D-E; Table 15; Pl. 16, Figs 3-8 . . . 43, 48, 54-55, 79, 84-85, 167, 257, 269; Text-figs 23, M. lindneri Glenister,1958...... 150 56A-F; Table 29; Pl. 27, Fig. 12, Pl. 33, Figs 1-2, 5-8 M. neverovi Bogoslovsky,1958...... 150 L. sp...... 167, 269; Pl. 33, Figs 3-4 M. nodifer (Clarke, 1885) ...... 27, linguum, Linguatornoceras ...... 167, 257; Pl. 27, Figs 13-14 79,83,123,238;Text-figs 23, 35B; Pl. 18, Figs 11-12 linguum, Linguatornoceras aff...... M. oxy Clarke,1897...... 55,137, 142 ...... 43,48,54-55,79 , 84-85, 167, 257, 269; Text-figs M. Pattersoni Clarke,1898...... 23, 56A-F; Table 29; Pl. 27, Fig. 12, Pl. 33, Figs 1-2, 5-8 . . . . 113, 115, 117, 119, 124-125, 127, 129-132, 221, 230 linguum, Tornoceras (Linguatornoceras)aff...... 166 M. pattersoni var. styliophilum Clarke, 1898 . . . . . 113, 115, 119 Lobotornoceras Schindewolf,1936...... M. regulare Fenton&Fenton,1924...... 129 ...... 65,73, 162, 168, 171-172; Text-fig. 21 M. rhynchostomum Clarke,1898...... 142 L. ausavense (Steininger, 1853) ...... M. schellwieni Wedekind,1913...... 137 ...... 168, 171-172; Text-figs 58C, F M. septentrionale Miller, 1938 ...... 129, 150 L. bicaniculatum Petter,1959...... 173 M. simulator (Hall, 1874) . . 83, 124, 130, 221; Pl.9,Figs4-5 L. hassoni House, 1978 ...... 172; Text-fig. 58E M. sinuosum (Hall, 1843) ...... HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 279

. . . . 1, 34, 38-42, 108, 115, 123, 129, 131-132, 135, 144 165, 258; Text-figs 23, 54G, 55C; Table 25; Pl. 28, Figs 6-10 M. cf. sinuosum House&Pedder,1963...... 129 orbiculus, Ammonites ...... 142 M. sinuosum apprimatum Clarke,1898...... orientale, Acanthoclymenia ...... 102 ...... 1,27,79,83,123, 132, 135, ovata, Tornoceras simplex ...... 167 185, 237; Text-figs 23, 35B, 42F, 63; Pl. 17, Figs 5-10 oxy, Carinoceras ...... 144 M. sinuosum clausium n.ssp...... 1, oxy, Manticoceras ...... 55,137, 142 35, 40-41, 79, 129-131, 226, 229; Text-figs 23, 39A-I, oxy, Sphaeromanticoceras ...... 40A-B; Table 12; Pl. 12, Figs 1-7, 11, Pl. 13, Figs 1-15 . . 1, 43, 45, 48, 54-55, 57-58, 79, 85, 135, 140, 143-144, 145, M. sinuosum sinuosum (Hall, 1843) ...... 35,38- 148; Text-figs 23, 42D, 44A, C, E-F, 45A-D, 46B; Table 19 42, 79,83,124-127, 129-130, 132, 135, 221-222, 225- oxy, Sphaeromanticoceras aff...... 43,48,85,140,148;Text-fig. 44F 226, 233-234; Text-figs 23, 36A-G, 37A-H, 38; Table 11; Oxytornoceras Becker,1993...... 73, 77, 86, 162, 169; Text-fig. 21 Pl.9,Figs6-8,Pl.10,Figs1-12,Pl.11,Figs1-13,Pl.12, O. acutum (Frech, 1902) ...... 169 Figs 8-10; Pl. 15, Figs 1-2, Pl. 16, Figs 11-12 paradoxum, Neomanticoceras ...... 217; Pl.7,Figs2-3 M. sinuosum tardum Clarke,1898...... Parodiceras Hyatt, 1884 ...... 162 ...... 1,40,79, 130, 132, 221, 230; Text-figs 23, ParodiceratinaePetter,1959...... 162 39J, 40C; Table 13; Pl. 9, Figs 2-3, Pl. 14, Figs 1-10 pattersoni, Goniatites ...... 108 M. tardum Clarke,1898...... 130-132 Pattersoni, Manticoceras ...... M. unduloconstrictum Miller, 1938 ...... 141 ...... 113,115,117,119,124-125,127,129-132,221,230 M. sp...... 137, 237; Pl. 17, Fig. 4 paucistriatum, Aulatornoceras ...... 54-55, 79, 85, 175-176, 178- manticoides, Chutoceras ...... 107-109 180, 266; Text-figs 23, 60D, F; Table 33; Pl. 32, Figs 2-13 Maternoceras Clausen,1971...... 73, 77, 122; Text-fig. 21 peracutum, Epitornoceras ...... 24,79, 169- M. sandbergeri (Wedekind, 1910)...... 85 170,171,185;Text-figs 23, 57A-E; Table 28; Pl. 34, Fig. 8 menneri, Carinoceras ...... 137-138 peracutum, Epitornoceras aff...... 16,169-171; Text-figs 57B-C Meropharciceras Becker & House, 1993 ...... 73, 76; Text-fig. 21 peracutum, Epitornoceras cf. . . . . 16, 170-171, 185; Text-figs 57D-E, 63 Mesobeloceras Glenister,1958...73, 77, 83, 155, 160-161; Text-fig. 21 perlatum, Ponticeras ...... 16-17, 20, milleri, Maeneceras ...... 3, 62, 185; Text-figs 61E, 63 24-26, 79,82,98-99, 100-102, 108, 185, 210, 213; Text-figs milleri, Sporadoceras ...... 62 28A-G, 29A-E, 63; Table 4; Pl. 4, Figs 5-9, Pl. 5, Figs 1-9 mithracoides, Epitornoceras ...... 82,169-170; Text-fig. 57G perlatum, Ponticeras cf...... 16,23,101-102; Text-figs 29D-E mithracoides, Epitornoceras cf. . . . 16, 24, 82, 170, 185; Text-figs 57F, 63 Petteroceras Bogoslovsky,1967...... 73, 76, 82; Text-fig. 21 mithracoides, Goniatites ...... 169 Petteroceratidae Becker & House, 1993 . . . 72-73, 89; Text-figs 20-21 Mixomanticoceras Becker et al.,1993...... 122; Text-fig. 21 Pharciceras Hyatt,1884...... Mzerrebites Becker&House,1993...... 73, 76; Text-fig. 21 ...... 3,10-11, 66, 73, 76, 89-92, 186, 205; Text-fig. 21 naplesense, Naplesites ...... P.amplexum (Hall, 1886) . . 10-11, 79, 82, 89-90, 91, 185-186, . . 79, 160-161, 186, 253; Text-figs 23, 52F-H; Pl. 25, Fig. 6 205; Text-figs 23, 26D-H, 63; Table 2; Pl. 1, Figs 1-8 naplesense, Neomanticoceras ...... 160 P.barnetti Work,Mason,&Klapper,2007...... 186 naplesense, Probeloceras? ...... 160, 253 P.bidentatum Petter,1959...... 89 Naplesites Yatskov,1991...... P. galeatum Wedekind, 1918 ...... 89, 170 ...4,73, 77, 83, 155-157, 160-161, 186, 253; Text-figs 21, 52 P. tridens (G. & F. Sandberger, 1850) ...... N. anguisellatum (Chao, 1956) ...... 160-161 ...... 1,89-91, 205; Text-figs 26 A-C; Pl. 1, Fig. 10 N. housei (Montesinos & Henn, 1986) ...... 161 P.? sp. . . 13, 27, 79, 92, 265; Text-fig. 23; Pl. 31, Figs 14-15 N. inyx (Clarke, 1898)...... 79, 160-161, PharciceratidaeHyatt,1900...... 72-73, 89; Text-figs 20-21 185-186, 253; Text-figs 23, 52A-E, 63; Pl. 25, Figs 2-5 Phoenixites Becker, 1993 ...... 60,73, 77, 173; Text-fig. 21 N. cf. inyx Becker, 1993 ...... 161 P. concentricus (House, 1965)...59-60, 79, 86, 158, 173, 261; N. naplesense (Clarke, 1898)...... 79, Text-figs 23, 54M-O, 56A; Table 26; Pl. 29, Figs 1-15 160-1671, 186, 253; Text-figs 23, 52F-H; Pl. 25, Fig. 6 P. sp...... 62 N. sp. (Bogoslovsky, et al., 1982) ...... 160 planorbe, Acanthoclymenia ...... 102 neapolitana, Acanthoclymenia ...... 35,39-40, 79, 83, 103- Playfordites Becker et al., 1993 . . . 73, 84-85, 122, 135, 141; Text-fig. 21 106, 214; Text-figs 23, 30E-H; Table 5; Pl. 6, Figs 1-15 P. tripartitus (G. & F. Sandberger, 1850) ...... 141 neapolitana,Acanthoclymeniaaff...... P. cf. tripartitus (G. & F. Sandberger, 1850)...... 31,33,79,83,102-105; Text-figs 23, 30D ...... 43,48,54,57,79, 84- neapolitana, Acanthoclymenia cf...... 106 85, 135, 142, 185; Text-figs 23, 42A, 63; Table 17 Neomanticoceras Schindewolf, 1936 . . 73, 84-85, 122, 160; Text-fig. 21 pompeckji, Maeneceras ...... 86 N. naplesense (Clarke, 1898) ...... 160 pompeckji, Maeneceras cf...... 62,86 N. paradoxum (Matern, 1931) ...... 217; Pl. 7, Figs 2-3 pompeckji, Sporadoceras cf...... 183 Neopharciceras Bogoslovsky 1955 ...... 73, 76-77, 82, 89; Text-fig. 21 pompeckji, Tornoceras ...... 166 neverovi, Manticoceras ...... 150 Ponticeras Matern, 1929 ...... 4,13,16,19- nitidum, Crassotornoceras ...... 168 21, 23, 27, 66, 73, 77, 82, 98, 100-103, 107-109; Text-fig. 21 nodifer, Manticoceras ...... P. aequabile (Beyrich, 1837)...... 98 27, 79, 83, 123, 238; Text-figs 23, 35B; Pl. 18, Figs 11-12 P. auritum (Holzapfel, 1899) ...... 98 nundaia, Goniatites ...... 107 P. barroisi (Wedekind, 1917,1918) ...... 98 nundaium, Chutoceras ...... 1, 17, 23, 25-26, 77, 79, 82, 100, P. bisulcatum (Keyserling, 1844)...... 98 107-109, 185, 221; Text-figs 23, 28H-K, 63; Pl. 9, Fig. 1 P. discoidale Glenister,1958...... 98 obseum, Tornoceras uniangulare ...... 35,39-41,79,83,158, P. domanicense (Holzapfel, 1899) ...... 98 280 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

P. kayseri (Wedekind, 1917, 1918) ...... 98 27G-H, 63; Table 3; Pl. 2, Figs 4-7, Pl. 3, Figs. 5-6 P. keyserlingi (Holzapfel, 1899) ...... 98 S.? syngonum Miller,1938...... 95 P. perlatum (Hall, 1874, 1876).....16-17, 20, 24-26, 79, 82, Sandbergeroceratidae Miller, 1938 ...... 89 98-99, 100-102, 108, 185, 210, 213; Text-figs 23, 28A-G, sandbergerorum, Sandbergeroceras ...... 92-95; Text-fig. 27F 29A-E, 63; Table 4; Pl. 4, Figs 5-9, Pl. 5, Figs 1-9 schellwieni, Manticoceras ...... 137 P. cf. perlatum (Hall, 1874)...... Schindewolfoceras Miller,1938.....49,73, 77, 84, 92, 96; Text-fig. 21 ...... 16,23,101-102; Text-figs 29D-E S. chemungense (Vanuxem, 1842)...... 50,79, 84, P. cf. regale (Holzapfel, 1899) ...... 82, 107-108 94, 96-97, 98, 206; Text-figs 23, 27I, 63; Pl. 2, Fig. 8 P. ? sp...... 31,79, 83, 102, 115; Text-figs 23, 33M S. aff. chemungense (Vanuxem, 1842)...... posterior, Aulatornoceras eifliense ...... 175 ...... 94,97, 210; Text-fig. 27J; Pl. 4, Figs 1-4 Posttornoceratidae Bogoslovsky, 1962 ...... 72, 161-162; Text-fig. 20 S.? equicostatum (Hall, 1874) ...... primordalis, Ammonites ...... 142 ...50,79, 84, 96-97, 98, 209; Text-fg. 23; Pl. 3, Fig. 4 Probeloceras Clarke,1898...... 4,35, S.? aff. equicostatum (Hall, 1874) ...... 73, 82, 102-103, 105, 109, 155-156, 157, 160; Text-fig. 21 ...... 50,79, 84, 96-98, 209; Text-fig. 23; Pl. 3, Fig. 1 P. alveolatum Glenister,1958...... 109 ?Schindewolfoceras sp...... 51,97, 206; Pl.2,Fig.3 P. cf. genundewa Clarke, 1898; House, 1978 ...... 106 Semicheiloceras Becker, 1993 ...... 162 P.lutheri (Clarke, 1855)...... septentrionale, Manticoceras ...... 129, 150 ...... 35,38-43,79, 95, 105, 125, 152, 155- Serramanticoceras Becker, House, & Kirchgasser, 1992 ...... 156, 157, 159-161, 185, 250, 253; Text-figs 23, 35C, 50- ...... 73, 77; Text-fig. 21 51, 63; Table 22; Pl. 24, Figs 5-9, Pl. 25, Fig. 1 serriense, Aulatornoceras ...... 176 P. lutheri n. ssp. Becker et al., 1993 ...... 157 simplex, Ammonites...... 163 P. aff. lutheri Becker et al.,1993...... 157 simplex, Tornoceras ...... 163 P. cf. lutheri Becker et al., 1993 ...... 157, 160 simulator, Goniatites ...... 122-124 P.strix Kirchgasser, 1975 ...... 83, 109 simulator, Manticoceras ...... 83,124, 130, 221; Pl.9,Figs4-5 P. aff. strix Kirchgasser, 1975 ...... 110 sinuosum, Manticoceras cf...... 129 P.? naplesense Clarke,1898...... 160, 253 sinuosum, Manticoceras ...... Prochorites Clausen, 1969 . . . . 73, 98, 102, 105, 109, 157; Text-fig. 21 ...... 1, 34, 38-42, 108, 115, 123, 129, 131-132, 135, 144 P.alveolatus (Glenister, 1958) ...... sinuosum sinuosum, Manticoceras ...... 35, 38-41, 79, 83, 109-110, 156-157, 185, ...... 35,38-42, 79, 83, 124-127, 129-130, 132, 217; Text-figs 23, 31, 32C, 63; Table 7; Pl. 7, Figs 4-6 135, 221, 225-226, 233-234; Text-figs 23, 36A-G, 37A-H, 38; P. aff. alveolatus (Glenister, 1958) . . . . 49, 79, 110, Text-fig. 23 Table 11; Pl. 9, Figs 6-8, Pl. 10, Figs. 1-12, Pl. 11, Figs 1-13, Prolecanites Majsisovics, 1882 ...... 97 Pl. 12, Figs 8-10, Pl. 15, Figs 1-2, Pl. 16, Figs 11-12 Pseudarietites Frech, 1902 ...... 92 sinuosus, Goniatites ...... 107-108, 124 Pseudoclymeniidae sensu Becker,1995...... sororium, Carinoceras ...... 72-73, 78, 161-162; Text-figs 20-21 ...79, 85, 137, 238; Text-figs 23, 42G; Pl. 18, Figs 3-4, 6-9 Pseudofordites Bogoslovsky, 1959 ...... 76 Sphaeromanticoceras Clausen,1971...... Pseudoprobeloceras Bensaïd,1974.....73, 76, 98, 120, 122; Text-fig. 21 57, 73, 77, 122, 129, 134-135, 137, 140, 142, 149; Text-fig. 21 Puncticeras Becker,1993...... 181 S. affine (Steininger, 1849) ...... 142, 157 Raymondiceras Schindewolf 1934 ...... 181 S. oxy (Clarke, 1897) ...... 1,43,45, regale, Ponticeras cf...... 82,107-108 48, 54-55, 57-58, 79, 85, 135, 140, 143-144, 145, 148; regulare, Manticoceras ...... 129 Text-figs 23, 42D, 44A, C, E, 45A-D, 46B; Table 19 rhynchostomum, Manticoceras ...... 142 .S. aff oxy (Clarke, 1897)...43,48,85,140, 148; Text-fig. 44F rhynchostomum, Sphaeromanticoceras ...... 1, 43, 48-49, S. rickardi House&Kirchgasser,1993...... 58-59, 79, 51, 54-57, 79, 84-85, 134-135, 139, 142-144, 148, 185, 242, 148-149, 155, 185; Text-figs 23, 48A-C, 63; Table 20 245-246; Text-figs 23, 41F, 42B, H-I, 43F-G, 47, 63; Table 18; S. aff. rickardi House&Kirchgasser,1993...... Pl. 20, Figs 1-12, Pl. 21, Figs 1-12, Pl. 22, Figs 1-9 ...... 58,79, 85, 149-150; Text-fig. 23 rhysum, Aulatornoceras ...... 58-59, 79, S. rhynchostomum (Clarke, 1898) . . 1, 43, 48-49, 51, 54-57, 79, 85, 175-176, 178-179, 266; Text-figs 23, 60E; Pl. 32, Fig. 1 84-85, 134-135, 139, 142-144, 148, 185, 242, 245-246; rickardi, Sphaeromanticoceras ...... 58- Text-figs 23, 41F, 42B, H-I, 43F-G, 47, 63; Table 18; Pl. 59, 79, 148-149, 155, 185; Text-figs 23, 48A-C, 63; Table 20 20, Figs 1-12, Pl. 21, Figs 1-12, Pl. 22, Figs 1-9 rickardi, Sphaeromanticoceras aff...... 58,79, 85, 149-150; Text-fig. 23 Sphaeropharciceras Bogoslovsky, 1955 ...... 89 sagittarium, Beloceras ...... 94;Text-fig. 27C Sporadoceras Hyatt, 1884 ...... 3, 73, 87, 182-183; Text-fig. 21 sandbergeri, Aulatornoceras ...... 175 S. milleri (Flower & Caster, 1935) ...... 62 sandbergeri, Maternoceras ...... 85 S. cf. pompeckji Wedekind, 1918 ...... 183 sandbergeri, Tornoceras ...... 179 Sporadoceratidae Miller & Furnish, 1957 ...... sandbergeri, Truyolsoceras ...... 179 ...... 72-73, 182-183; Text-figs 20-21 Sandbergeroceras Hyatt, 1884 . . . 49-50, 73, 83, 92-93, 95; Text-fig. 21 Staffites Wedekind,1918...... 77,181 S. Chemungense Clarke1898...... 97 Stenopharciceras Montesinos & Henn, 1986 . . . 73, 76, 89; Text-fig. 21 S.? enfieldense n.sp...... 1,49-51, 79, 95, strix, Eidoprobeloceras ...... 109 206, 209; Text-fig. 23; Pl. 2, Figs 1-2, Pl. 3, Figs 2-3 strix, Probeloceras ...... 83,109 S. sandbergerorum Miller, 1938 ...... 92-95; Text-fig. 27F strix, Probeloceras aff...... 110 S. syngonum (Clarke, 1897) ...... styliophilum, Manticoceras pattersoni var...... 113, 115, 119 1, 34, 42, 49-50, 79, 93-95, 185, 206, 209; Text-figs 23, styliophilus, Koenenites ...... 22,26,112 HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 281

styliophilus, Koenenites aff...... 119 T. sp. House, 1965 ...... Text-fig. 54P styliophilus styliophilus, Koenenites ...... 1,17, T.? n.sp.House,1965...... 257; Pl. 27, Fig. 11 19-20, 22-23, 79, 82, 112-113, 115-117, 119-120, 185, 218; Tornoceratidae Arthaber, 1911 ...... Text-figs 23, 34A-F, I-J, L, 35A, 63; Table 8; Pl. 8, Figs 1-9 ...... 71-72, 73, 161-162, 172-173, 175; Text-figs 20-21 styliophilus styliophilus morphotype B, Koenenites ...... 115,117,120 TornoceratinaeArthaber,1911...... 162 subpartitus, Goniatites ...... 181 Tornoceratini Arthaber, 1911 ...... 162, 172-173 syngonum, Sandberberoceras ...... Trevonites Becker & House, 1994 ...... 73, 162, Text-fig. 21 ...... 1,34,42,49-50,79, 93-95, 185, 206, 209; Text- TriainoceraeHyatt,1884...... 89,92 figs 23, 27G-H, 63; Table 3; Pl. 2, Figs 4-7, Pl. 3, Figs 5-6 Triainoceras Hyatt,1884...... 91-93 syngonum, Sandbergeroceras? ...... 95 T. costatum (d’Archiac & de Verneuil, 1842) ...... Synpharciceras Schindewolf, 1940 ...... 73, 76, 89; Text-fig. 21 ...... 94;Text-fig. 27E Synpharciceratidae Schindewolf, 1940 ...... 89 T. gerassimovi Bogoslovsky,1958...... 92,94;Text-fig. 27D Tamarites Bogoslovsky, 1965 ...... 73, 91-92; Text-fig. 21 T. incertum (d’Archiac & de Verneuil, 1842) ...... 92 tardum, Manticoceras ...... 130-132 T.? tuberculosum (d’Archiac & de Verneuil, 1842) ...... 92 tardum, Manticoceras sinuosum ....1,40,79, 130, 132, 221, 230; Text- Triainoceratidae Hyatt, 1884 ...... figs 23, 39J, 40C; Table 13; Pl. 9, Figs 2-3, Pl. 14, Figs 1-10 1, 50, 63, 71-72, 73, 83-84, 88-89, 91-92, 94; Text-figs 20-21 Timanites Mojsisovics, 1882 ...... 73, 83, 112; Text-fig. 21 tridens, Pharciceras ....1,89 -91, 205; Text-figs 26 A-C; Pl. 1, Fig. 10 Timanoceras Bogoslovsky,1957...... 73, 77, 122; Text-fig. 21 Trimanticoceras House in House & Ziegler, 1977 ...... Tornia House,1970...... 73, 78, 162 ...... 73, 77, 122; Text-fig. 21 Tornoceras Hyatt,1884...... 11,16,19-21, 27-34, 41-42, 60, 65, T. cinctum (Glenister, 1958) ...... 85 73, 76, 157-158, 162-167, 169, 171, 173; Text-figs 21, 53-54 tripartitus, Playfordites ...... 141 T. acutum Frech, 1902 ...... 169 tripartitus, Playfordites cf...... 43,48,54, T. arcuatum House, 1965 . . 22, 79, 82, 158, 165-166, 170; Text- 57, 79, 84-85, 135, 142, 185; Text-figs 23, 42A, 63; Table 17 figs 23, 54H-J, L; Pl. 26, Figs 7-10, Pl. 27, Fig. 16 tripartitus, Goniatites lamed var...... 141 .T. cf arcuatum House,1965...... Truyolsoceras Montesinos, 1987 ...... 73, 175, 179; Text-fig. 21 ...... 10,158, 166, 170; Text-fig. 54K T. bicostatum (Hall, 1843)...... 1,59- T. ausavense crassum Matern1931...... 168 61, 79, 86, 175-176, 178-179, 180, 257, 262; Text-figs 23, T. belgicum (Matern, 1931) ...... 168 59B, 60A-C, H; Table 34; Pl. 30, Figs 2, 4-13, 15 T. bicostatum (Hall, 1843) ...... 178 T. cf. bicostatum (Hall, 1843) ...... 59-61, 79, 86, 180, T. (Aulatornoceras) bicostatum (Hall, 1843) ...... 178-179 262, 265; Text-fig. 23; Pl. 30, Fig. 15, Pl. 31, Fig. 1 T. (Aulatornoceras) clarkei Miller, 1938 ...... 61 T. clarkei (Miller, 1938)...... 59,61, T. clausum Glenister, 1958 ...... 166 79, 86, 176, 179-180, 262; Text-fig. 23; Pl. 30, Figs 1, 3 T. (Tornoceras) concentricum House,1965...... 173 T. sandbergeri (Ford & Crick, 1897)...... 179 T. Edwin-halli Clarke,1898...... 62,87,183 tuberculoso-costatus (pro, pars) G. & F. Sandberger, 1850, Goniatites ...93 T. eifliense (Steininger, 1849)...... 177 tuberculosum, Trianoceras? ...... 92 T. frechi Wedekind, 1918 ...... 173 tynani, Wellsites ...... 50-51, 79, T. guestphalicum Frech, 1897 ...... 166 83-84, 94, 96, 185, 205; Text-figs 23, 27A, 63; Pl. 1, Fig. 9 T. haugi Frech,1902...... 166 typum, Tornoceras ...... 16,163 T. (Linguatornoceras)aff. linguum House, 1965 ...... 166 typum, Tornoceras cf...... 43,48- T. pompeckji Wedekind 1918 ...... 166 49, 55, 85, 157, 166, 257; Text-figs 53C-D; Pl. 27, Fig. 15 T. sandbergeri (Ford & Crick, 1897)...... 179 typus, Goniatites retrorsus var...... 163 T. simplex (von Buch, 1832)...... 163 Uchtites Bogoslovsky,1969...... 73, 98; Text-fig. 21 T. simplex mut. ovata (Münster, 1832) Frech, 1902 ...... 167 undulatus, Goniatites ...... 175 T. typum (G. & F. Sandberger, 1851) ...... 16,163 unduloconstrictum, Delphiceras ...... 238; Pl. 18, Fig. 10 T. cf.typum(G. and F. Sandberger, 1851) ...... 43,48-49, unduloconstrictum, Manticoceras ...... 141 55, 85, 157, 166, 257; Text-figs 53C-D; Pl. 27, Fig. 15 uniangulare, Tornoceras ...... Text-fig. 23 T. uniangulare (Conrad, 1842) ...... uniangulare, Tornoceras aff...... Text-fig. 23; Pl. 27, Figs 1-6 ...... 39,163, 165-166, 169; Text-fig. 23 uniangulare, Tornoceras cf...... T. aff. uniangulare (Conrad, 1842) ...... 79,158, 164, 185; Text-figs 23, 54D-E, 63 ...11,79, 158, 164, 257; Text-fig. 23; Pl. 27, Figs 1-6 uniangulare uniangulare, Tornoceras . . . 1, 10-12, 79, 158, 160, 163, 165, T. cf. uniangulare (Conrad, 1842) ...... 254; Text-figs 23, 54A-C, 55A; Table 23; Pl. 26, Figs 1-6 ...... 79,158, 164, 185; Text-figs 23, 54D-E, 63 uniangulare widderi, Tornoceras ...... 81,254; Pl. 26, Fig. 5 T. uniangulare arkonensis (House, 1965) ...... 81 vagans, Carinoceras ...... 43,48,54-55, 79, 84-85, 138-140, T. uniangulare compressum Clarke,1897...... 249; Text-figs 43A-E, 44D, 46A; Table 16; Pl. 23, Figs 5-16 ...... 27,29,79, 82, 157-158, 162, 164-165, 158; Text- vagans, Carinoceras aff...... figs 23, 53A-B, 54F, 55B; Table 24; Pl. 28, Figs 1-5 ...... 55,79, 138, 141, 249; Text-fig. 23; Pl. 23, Figs 1-4 T. uniangulare obesum Clarke,1897...... vericosum, Archoceras ...... 85 ...... 35, 39-41, 79, 83, 158, 165, 258; Virginoceras Ruzhencev,1960...... 73, 122; Text-fig. 21 Text-figs 23, 54G, 55C; Table 25; Pl. 28, Figs 6-10 Virginoceratinae Yatskov, 1990 ...... 122 T. uniangulare uniangulare (Conrad, 1842) ...... wabashense, Archoceras ...... 237; Pl. 17, Fig. 1 ...... 1, 10-12, 79, 158, 160, 163, 165, 254; Wedekindella Schindewolf, 1928 ...... 73, 76, 162; Text-fig. 21 Text-figs 23, 54A-C, 55A; Table 23; Pl. 26, Figs 1-4, 6 Wellsites House & Kirchgasser, 1993 ...... T. uniangulare widderi House,1965...81, 254; Pl. 26, Fig. 5 ...... 49,73,77,79, 84, 92, 95-96; Text-fig. 21 282 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

W. tynani House & Kirchgasser, 1993 ...... 50-51, 79, 83-84, Werneroceras Wedekind, 1918 ...... 76, 162 94, 96, 185, 205; Text-figs 23, 27A, 63; Pl. 1, Fig. 9 widderi, Tornoceras uniangulare ...... 81, 254; Pl. 26, Fig. 5 W. williamsi (Wells, 1956) [Beloceras]...... williamsi, Wellsites ...... 50,79, 94, 96; Text-figs 23, 27B ...... 50,79, 94-96; Text-figs 23, 27B

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 bold font.

AbbeyGulfBeds(PennYanShale)...... 16,20 15, Pl. 9, Figs 2-3, 6-8, Pl. 10, Figs 1-4, 10, 12, Pl. 12, Figs 1- Angola Shale [marker (Scraggy Bed) black shale at base; top Rhinestreet 7, 11, Pl. 13, Figs 1-15, Pl. 14, Figs 1-10, Pl. 16, Fig. 12, Pl. 24, Shale]...... 48-49, 54-55, 57; Text-figs 3-4, 13, 15 Figs 6, 8-9, Pl. 25, Fig. 1, Pl. 31, Figs 7-17 Angola Shale (microcycles/minor rhythms abca; 1-9 of lower part) . .... Cashaqua Shale (upper part or division of Kirchgasser, 1975) ...... 51-52, 65, 85; Text-figs 15, 19 ...... 36-37, 68, 105-106, 109-110, Angola Shale (West Falls Group) . . . . 4-5, 6, 43, 45, 48-49, 51-52, 54- 157, 222, 225-226, 233-234, 258, 265; Text-figs 10, 30E-H, 55, 64-65, 68-69, 79-80, 84-85, 123, 134-144, 148, 157, 166- 31A-H, 32, 38, 54-55; Pl.6,Figs1-15,Pl.7,Figs4-6,Pl.10, 169, 175-180, 184, 234, 238, 242, 245-246, 249, 265-266, 269; Figs 5-9, 11, Pl. 11, Figs 1-13, Pl. 12, Figs 8-10, Pl. 15, Figs 1- Text-figs 2-3, 4, 15, 23-24, 41-47, 53, 60, 63; Pl. 16, Figs 1-10, 2, Pl. 16, Fig. 11, Pl. 28, Figs 6-10, Pl. 31, Figs 18-20 Pl. 18, Figs 3-4, 6-9, Pl. 20, Figs 1-2, 9-12, Pl. 21, Figs 1-12, Cashaqua Shale/Rhinestreet Shale contact (Sonyea/West Falls Groups) . . Pl. 22, Figs 1-8, Pl. 23, Figs 1-4, 7-12, 15-16, Pl. 31, Figs 2-3, ...... 5, 34, 36-37; Text-figs. 3-4, 10 5-6, Pl. 32, Figs 2-14, Pl. 33, Figs 1-4, 9-20 Catskill (Delta, clastic sequences; facies; red-bed succession; marine/non- Angola Shale equivalents ...... marine)...... 1-3, 63-64, 84; Text-fig. 18 ...... 48-49, 51, 57, 68; Text-figs 2-3,4,15, 41 Cattaraugus Shale and Sandstone (Conewango Group) ...... ApuliaMember(LowerTullyFormation)...... 11;Text-fig. 6 ...... 4,80, 87, 183-184; Text-figs 2, 24, 63 Arkona Shale (Ontario, Canada) ...... 81 Cayuta Shale, Upper (Chesney Formation; Member, Chemung Group/ AssisedeFrasnes,lower(Europe)...... 70 Formation;RhinestreetShaleequivalent)...... Assise de Matagne (Belgium) ...... 168 ...... 50, 96-98, 206, 209; Pl.2,Fig.8,Pl.3,Fig.1 BeardsCreekHorizon(BC;WestRiverShale)...... CenterfieldLimestone(HamiltonGroup)...... 81-82 31-33, 35, 79, 102, 114-117, 122; Text-figs 9, 23, 33-34 ChadakoinShale(EllicottShale,ConeautGroup)...... BedG,WestBrookMember(UpperTullyFormation)...... 4,40,80, 86, 183-184; Text-figs 2, 24, 63 ...... 10-11; Text-fig. 6 ChagrinMember(ConneautGroup)...... 62 BedfordShale(ConewangoGroup)...... 62,87 ChemungGroup(Formation;rocks)...... Beers Hill Member (Rhinestreet Shale) ...... 50-51 ...... 5, 26, 51, 98, 107, 140; Text-figs 3-4; Pl. 2, Fig. 3 BelgianF2dreefs(Belgium)...... 70 Chemung Group (Grimes Sandstone at base; Naples, Ontario County) . BelgianF2hreefs(Belgium)...... 70 ...... 5, 140; Text-fig. 3 Bellona Coral Bed (Tully Formation)....10-11, 66, 82; Text-figs 5-6 ChemungGroup,Lower...... 26,98,107 Bergen Beach Bed (Lower Hubbard Quarry Submember Geneseo Shale) CherryValleyLimestone(MarcellusFormtion,HamiltonGroup)...... 23 ...... 81,161-162 Bituminous(Black)shale(ofClarke,1904;RhinestreetShale)...... 43 Chittenango Shale (Marcellus Formation, Hamilton Group) . . . 81, 163 BlackShaleandSlate(GeneseeGroup)...... 12 ClevelandShale(Ohio;ConewangoGroup)...62,87,183;Text-fig.61F Bluestone Bed (Ithaca Shale and Sandstone) (Genesee Group) . . . . 17, 26 Conewango Group ...... 3-4, 9, 62, 183; Text-figs 2, 61E Bluff Point Silstone (BP; West River Shale) ...... 30-34, 65, 79,83, ConneautGroup(Formation)...... 103, 114, 117, 121, 218; Text-figs 23, 33-34; Pl. 8, Figs 10-11, ...... 4,9,62, 183, 273; Text-fig. 2; Pl. 35, Figs 11-13 14Conodont Bed (of Hinde, 1879; = North Evans Ls; Genesee Group)... Brallier Shale (or equivalents; Pennsylvania, West Virginia, Virginia)...... 13,27 ...... 157 Corning Shale (Member, Rhinestreet Shale) ...... 4, 49; Text-fig. 2 Büdesheimer Schiefer (Germany) ...... 167-168, Corell’sPointGoniatiteBed(CP;GowandaShale)...... 172, 217, 257; Text-fig. 58; Pl.7,Figs2-3, Pl. 27, Figs 13-14 ...... 5, 60, 79, 158, 174-175, 182, 257; Text- Burlington Limestone (Missouri) . . . 94; Text-fig. 27; Pl. 4, Figs 1-4 figs 3, 17, 23, 54M-O, 59B-C, 60A-C, 61A-B; Pl. 27, Figs 7- Callaway Limestone (Missouri) ...... 94 10, Pl. 29, Figs 1-15, Pl. 30, Figs 2, 4-14, Pl. 35, Figs 1-10 Canadaway Group ...... CrosbySandstone(CS;PennYanShale)...... 12,17, 21-25, 28, 30, ...... 4,9,57,59, 161-163, 173, 179-180, 182; Text-fig. 2 65, 79, 82, 103, 107, 113, 115, 117, 119, 165-166, 218, 254; Cashaqua Shale (marker black shales A-F) . . 35-37, 38-41; Text-fig. 10 Text-figs 7, 23, 34F, J, 54L; Pl.8,Figs6,9,Pl.26, Figs 7, 10 CashaquaShale(SonyeaGroup)...... Dasberg(unitinUpperDevonianofGermany)...... 77 ...... 1,3-5,6,34-35, 36-37, 38-42, 48-50, 64, 67-68, 79- DomanikSuite,Lower(Group;Timan,Russia)...... 70,108-109 80, 83, 93-95, 103-106, 108-110, 115, 125-127, 129-132, 152- Dunkirk Shale (Black Shale; Canadaway Group) . . 1, 4, 5,57,59,63-64, 154, 157, 165, 177, 184, 186, 206, 214, 217, 221-222, 225-226, 68, 70-71, 79-80, 86,149, 184; Text-figs 2-3, 16, 23-24, 63 229-230, 233-234, 250, 253, 265; Text-figs 2-3,4,10, 19, 23- Dunn Hill (Member, Rhinestreet Shale) ...... 4, 49, 51; Text-fig. 2 24, 27H, 30E-H, 31A-H, 35, 36A-G, 37A-H, 38, 39A-J, 40, EllicottShale(ChadakoinShale,ConeautGroup)...... 4,62,79,86, 50A-Q, 51A-E, 54G, 55, 63; Pl.2,Figs1-2,4-6,Pl.6,Figs1- 182-183, 273; Text-figs 2, 23, 61C-D, 62 ; Pl.35, Figs 11-13 HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 283

EnfieldShale(Formation,SonyeaGroup)...... 184, 241, 265; Text-figs 2-3,4,16, 23-24, 41G, 48C, G, 49A- ...4-5, 43, 50, 65, 95, 109; Text-figs 2-3; Pl. 3, Figs 2-3, 5-6 E, 60E, 63; Pl. 19, Figs 1-9, Pl. 31, Fig. 3, Pl. 32, Fig. 1 FarnhamCreekBed(turbiditeunit,lowerAngolaShale)...... Hanover Shale/Dunkirk Shale contact (West Falls Group/Canadaway ...... 51-52, 54-55, 65; Text-fig. 15 Group)...... 5-6, 56, 58-59, 86; Text-figs. 3-4, 16 FilmoreGlenBed(s)(Member,TullyLimestone)...... HanoverShaleequivalents(WestFallsGroup)...... 9-10, 11, 82, 92; Text-figs 5-6 ...... 5-6, 58-59; Text-figs 3-4, 16 FirTreeLimestone(Pyrite;Submember,GeneseoShale)...... Harrell Shale (Pennsylvania, West Virginia) ...... 12-13, 16, 23, 27-28, 102, 186; Text-fig. 7 . . . 68, 106, 112-113, 117, 120, 170-172; Text-figs 57B-C, 58E Firestone Bed(s) (facies; of Williams, 1884; Ithaca Shale and Sandstone) . Hatch Shale (Rhinestreet equivalent, West Falls Group) ...... 26, 108 ...... 5-6, 44-45; Text-figs 3-4, 11 Fossil Log Horizon (FL; Williamsburgh Bed; West River Shale) . . . . 31- Hemberg(unitinUpperDevonianofGermany)...... 77 33, 67, 79, 102-105, 122, 136; Text-figs 9, 23, 30D, 33K-L Hiatus concretion horizon (of Baird, 1976; Penn Yan Shale) . . 17, 21, 65 FossilLogHorizons...... 38,40,67, 79, 110; Text-fig. 23 High Point Sandstone (= Nunda Sandstone at Naples, Ontario County). Gardeau Shale (and Sandstone; Rhinestreet equivalent in part; West Falls ...... 140 Group) . . 4-5, 45, 48, 50, 65, 84, 148; Text-figs 2-3,4,12, 14 HighPointSandstone(WestFallsGroup)...... 95, 140 Genesee Group (Formation)...... 1,4-5, Hubbard Quarry Shale (Submember, Geneseo Shale)...... 7, 12-13, 14-15, 17, 19-20, 33-35, 64-66, 69, 79, 99, 102-103, ...... 13,16,23-24, 169, 171; Text-fig. 57E 105-109, 112, 114-116, 119, 124, 130, 136, 158, 163-164, 166, HuddleRiderBed(blackshaleinupperHanoverShale)...... 170-172, 186, 221; Text-figs 2-3,4,7-9, 14, 23,28A-K,32, ...... 56,59,71,86;Text-fig. 16 33A-O, 34, 54A-C, E-F, 57A, D-F; Pl. 9, Figs 1, 4-5 HungryHollowFormation(Ontario,Canada)...... 81-82 Genesee Slate ...... 12, 34 IthacaBeds(ofChemungGroup)...... 107 Geneseo Limestone Horizon (Geneseo Shale) ...... IthacaShaleandSandstone(Formation,GeneseeGroup)...... 1,13-15, 16, 18-20, 102; Text-fig. 8 ...... 4-5,6,12, 16-17, 21-24, 26-28, 30, 34, 65, 82-83, Geneseo Shale (black shale; Member, Genesee Group) ...... 100, 107-109, 113, 115, 119, 121, 170-171, 186; Text-figs 2-3, ...... 1,4-5,6,9-10, 12-15, 16, 18- 4, 28H-K, 33I-J, 57A, C; Pl.9,Figs1,4-5,Pl.34,Fig.8 21, 23-24, 27, 63-66, 79-80, 82, 92, 99, 102, 164, 169-171, JavaFormation(WestFallsGroup)...... 4,57;Text-fig. 2 184, 186, 210, 265, 270; Text-figs 2-3,4-7, 8, 23-24, 54E, 57D- Kellwasserkalk (Lower,Upper;Germany)...... 70-71, 85-86 F, 63; Pl. 4, Fig. 6, Pl. 31, Figs 14-15, Pl. 34, Figs 6-7 Knollenkalk facies...... 5, 35, 55, 57, 59, 66-67; Text-fig. 3 Geneseo (Shale)-Penn Yan (Shale) contact...... 5, 12-16, 18- Kramenzelkalk facies...... 66-67 21, 23-24, 79-80, 102, 171, 184; Text-figs 2-3, 7-8, 23-24, 63 Laona Sandstone (Siltstone; Canadaway Group)...... Genundewa Limestone (Genesee Group) ...... 4-5, 6, 59, 61, 86, 180; Text-figs 2-3,4; Pl. 30, Figs 1-3 ...... 1,4-5, 12-13, 14-15, 17-22, 27-31, 66-68, 70, 79- Leicester Pyrite (base of Geneseo Shale, Genesee Group) ...... 3-5, 80, 82-83, 91, 103-107, 112-113, 115, 117, 119-121, 123-124, 9-10, 12-13, 14-15, 16-20, 69, 81-82, 158-160, 163-164, 186, 130, 132, 135-136, 157-158, 162-165, 172, 184, 186, 218, 237- 254; Text-figs 2-3,4,7-8, 54A-C, 55A; Pl. 26, Figs 1-4, 6 238, 241, 250, 258; Text-figs 2-3,4,7-9, 23-24, 30A-C, I-L, Linden Horizon (LH; Linden Goniatite Horizon; Penn Yan Shale) . . . . . 33F-G, I-J, N-O, 34G, 35, 42F, 53A-B, 54F, 55, 63; Pl. 8, Figs ...... 14-15, 17-18, 19-23, 26, 79, 82, 103, 107, 115-116, 4-5, 7-8, 12-13, Pl. 17, Figs 5-10, Pl. 18, Figs 1-2, 5, 11-12, Pl. 119, 164, 186; Text-figs 8, 23, 34A, C-D; Pl. 8, Figs 3-4 19, Figs 10-16, Pl. 24, Figs 1-4, Pl. 28, Figs 1-5 LodiLimestone(LO)...... 5-6, 12-13, 14-16, 18-21, 23, 24-25, 28, Genundewa Limestone (Lower division) ...... 66, 79-80, 82, 99-102, 164, 184, 186; Text-figs 3-4, 7-8, 23- ...... 13,14-15, 17, 27-30, 33, 82, 106, 24, 28B-E, G, 29A-E, 32, 63; Pl.4,Figs5,7,Pl.5, Figs 1-9 112-113, 115, 117, 119-121, 123-124, 164, 218; Text-figs 8-9 Lodi Limestone beds A-D . . 16, 23-24, 28, 101-102; Text-fig. 29A-E Genundewa Limestone (Upper division) ...... LowerBlackBand(MiddlesexShale)...... 34,95 . 13-14, 15, 17-18, 27-30, 33, 83, 123-124, 136; Text-figs 8-9 LyaiolFormation(Timan,Russia)...... 176 Genundewa Limestone disconformity (sub-Upper Division)...... Marcellus Formation (Hamilton Group) ...... 162-163 ...... 12-15, 18, 27; Text-figs 7-8 MiddlesexShale(BlackShale;SonyeaGroup).....1,4-5, 6, 12, 17, 30- Genundewa Pyrite (Horizon) (pyritic nodular bed; Geundewa Limestone 34, 35-36, 37, 40, 42, 49-50, 63-64, 67, 69-70, 79-80, 83-84, equivalent) ...... 12, 30, 105, 107, 120-121; Text-fig. 7 92-93, 95, 102, 105, 115, 117, 121-122, 136, 184, 206; Text- Goniatite Concretionary Layer (Parrish Limestone, Cashaqua Shale)... figs 2-3,4,9-10, 23-24, 63; Pl.2,Fig.7,Pl.3,Figs5-6 ...... 42 MiddlesexShaleequivalents(SonyeaGroup)...... GogoFormation(WesternAustralia)...... 109, 157 ...... 33-34, 49-50, 83, 93, 95, 206; Pl. 2, Fig. 7 Gorge Gully Submember (Windom Shale)...... 11 MillboroShale(Virginia)...... 157 GowandaShale(CanadawayGroup)...... 3-5, 6, 59-60, 61-62, MillportMember(RhinestreetShale)...... 51 68, 71, 80, 86, 158, 161-163, 173, 185-176, 179-182, 184, 261- MontourShale[Lowertongue(Member)ofMiddlesexShale]...... 262; Text-figs 2-3,4,17, 24, 54M-O, 59A-C, 60A-C, 61A-B, ...... 4,34,49;Text-fig. 2 63; Pl. 29, Figs 1-15, Pl. 30, Figs 1-15, Pl. 31, Fig. 1 Moravia Bed (Bed G, West Brook Member, Tully Formation) ...... Grimes sandstone member (of Chemung formation of G. Chadwick) ...... 9-11, 66, 82, 91, 205; Text-figs 5-6; Pl. 1, Figs 1-8 ...... 140 MorelandShale(LowerMember,RhinestreetShale)...... Grimes Siltstone (Rhinestreet equivalent; West Falls Group) ...... 4,49-51, 84, 96, 98, 205; Text-figs 2, 27A; Pl. 1, Fig. 9 ...... 5-6, 45, 140, 249; Text-fig. 3; Pl. 23, Figs 13-14 MoscowShale(Formation,HamiltonGroup)...... 4-5, 10-11, 12-13, HamiltonGroup...... 17-20, 79-80, 164, 184,284;Text-figs 2-3, 6-7,8,23-24, 63 ...... 4-5, 12-13, 17, 158, 161-164, 169; Text-figs 2-3, 54D Moscow-Geneseo(Hamilton/Genesee)contact...... HanoverShale(WestFallsGroup)...... 3-5,6,56-57, 58-59, 67, ...... 4-5, 10, 12, 14-15, 18-21; Text-figs 2-3,4,7-8 70, 79-80, 85-86, 134, 141, 149-152, 155, 175-176, 178-180, Naples Group, Beds (beds, facies, Formation) ...... 284 BULLETINS OF AMERICAN PALEONTOLOGY no. 374

.....5, 34, 64, 83, 95, 110, 115, 123, 125-127, 132, 158, 161, Pulteney/Rock Stream turbidite wedge (Cashaqua Shale equivalents) . . . 166, 218, 221, 225, 237, 246; Text-figs 3,31A,34A, 36A-C, ...... 4-5, 6, 34-37; Text-figs 2-3,4,10 54G; Pl. 8, Figs 1-2,Pl.9, Figs 2-3, Pl. 11, Figs 6, 12-13, Pl. 14, Pulteney Shale (Member, Cashaqua Shale) . . . . 4-5,6,34;Text-figs 3-4 Fig. 3, Pl. 17, Fig. 5, Pl. 22, Figs 4-5, Pl. 27, Fig. 15 QuarrySandstoneinterval(IthacaFormation)...... 17, 26, 109 Nehden (Nehdener Schichten/Schiefer; -Schurbusch; Germany) ...... RelyeaCreekHorizon(RL)(upperRhinestreetShale)...... 3,71,77,181 ...... 43, 46-49, 79, 84, 134, 140-141, 143-144, 148, NewAlbanyShale(Indiana)...... 237; Pl. 17, Fig. 1, Pl. 18, Fig. 10 166-167; Text-figs 13, 23, 41F, 44F, 45B; Pl. 23, Figs 5-6 NewAlbanyShale(TrousdaleMember,Kentucky)...... 186 Renwick Shale (RE; Black Shale; Formation, Genesee Group) . . . 1, 4-5, North Evans Limestone (Conodont Bed of Hinde, 1879; Genesee Group) 6, 12, 16-18, 20-26, 79, 82, 102, 108, 186; Text-figs 2-3,4,23 ...... 12-15, 17-18, 27-28, 64; Text-figs 7-8 RhinestreetShale(BlackShale;WestFallsGroup)...... NundaSandstone(andequivalents;WestFallsGroup)...... 4-5, 6, 34-35, 38, 40, 43-47, 48-50, ...... 4-5, 6, 45, 51-53, 55-57, 55, 57, 64, 66-67, 69-70, 79-80, 83-84, 94-98, 110, 134-136, 59, 68, 108, 140, 143, 145, 148; Text-figs 2-3,4,15, 45C-D 138-140, 142-145, 148, 156, 161, 166-167, 176, 184, 186, 205- OldRedSandstone(England;Europe) ...... 3 206, 209, 242, 246, 249, 253, 257, 265, 269; Text-figs 2-3,4, OswayoShaleandSandstone(ConewangoGroup)...... 10-11, 13, 15, 23-24, 27A-B, I, 41B, F, 42A-B, E, 43D-E, 44F, ...... 4,80, 183-185; Text-figs 2, 24, 61E, 63 45B, 46-47, 52A-H, 56A-D, 63; Pl.1,Fig.9,Pl.3,Fig.1,Pl. Panama Conglomerate (Conewango Group)...... 4,62 20, Figs 7-8, Pl. 22, Fig. 9, Pl. 23, Figs 5-6, 13-14, Pl. 25, Figs Parrish Limestone (PL; Cashaqua Shale) ...... 2-5, Pl. 27, Fig. 12, Pl. 31, Fig. 4, Pl. 33, Figs 5-8 ...5-6, 35-37, 41-42, 43, 68, 79, 125-127, 129-130, 152-153, RhinestreetShale(microcylesofupperpart)...... 155, 157, 222; Text-figs 3-4, 10, 23, 36D-G, 37D, G-H, 50H, ...... 43-44, 46-47, 69; Text-fig.13 K, M-N, P, 51E; Pl. 10, Figs 1-2, 12, Pl. 24, Figs 5-9 RhinestreetShale/AngolaShalecontact...... PennYanBlackShaleA...... 14-15, 18-19, 20, 40; Text-fig. 8 ...... 5-6, 46-47, 52-53; Text-figs 3-4, 13, 15 PennYanBlackShaleB...... 14-15, 18-19; Text-fig. 8 RhinestreetShaleequivalents(WestFallsGroup)...... PennYanShale(GeneseeGroup)...... 4-5, . 5-6, 44-45, 46-47, 49-40, 57, 84, 97-98; Text-figs 3-4, 11-13 6, 12-13, 14-15, 16-24, 26-29, 33, 65, 68, 79-80, 82, 100-103, RockStreamSiltstone(SonyeaGroup)...... 107-109, 112-113, 115, 117, 119-120, 123, 164, 166, 170-171, ...... 4-5, 6, 34-37, 40-43, 68; Text-figs 2-3,4,10 184, 218, 270; Text-figs 2-3,4,7-9, 23-24, 28F, 34A-F, J, 35A, RockStreamSilstone/CashaquaShalecontact...... 57D, 63; Pl. 8, Figs 1-4, 6, 9, Pl. 34, Figs 6-7 ...... 5,36-37, 41-42; Text-figs 3, 10 PennYan(Shale)-Crosby(Sandstone)contact...... RoricksGlenShale(Member,RhinestreetShale)...... 4;Text-fig.2 ...... 12, 17, 22-23, 65; Text-fig. 7 RyePointShale(Member,CashaquaShale)...... 4,35;Text-fig. 2 PennYan(Shale)-Ithaca(ShaleandSandstone)contact...... SawmillCreekShale(UpperMember,MiddlesexShale).....34,49-50 ...... 5-6, 12, 22-23, 65; Text-figs 3-4, 7 SalternCoveGoniatiteBed(Devon,England)...167; Text-fig. 56E-F Pentonwarra Goniatite Bed (North Cornwall, England)...... 76 SB(Schumacher)blackshale(PennYanShale)...... Perrysburg Formation (Canadaway Group)...... 4-5; Text-fig. 3 ...... 14-16, 17-21, 102; Text-fig. 8 Pharciceras Bed(WestBrookMember,TullyLimestone)...... SchistesdeMatagne(MatagneShale;F3;Belgium)...... 10-11, 158, 205; Text-figs 6, 54K; Pl. 1, Figs 4, 7 ...... 70,140, 149-150, 168; Text-fig. 44B, 48B Pharciceras horizons (Trousdale Member, New Albany Shale, Kentucky) . SchumacherBed(SB;concretionbed,PennYanShale)...... 186 ...... 14-16, 18; Text-fig. 8 Pharciceras-Lage [Untere; Martenberg section, Diemelsee (Adorf), Ger- Scraggy Bed (concretion bed; marker black shale; top of Rhinestreet Shale) many]...... 91;Text-fig. 26A-C ...... 5-6, 43-44, 46-47, 48, 54, 148; Text-figs 3-4, 13, 15 PipeCreekShale(BlackShale)(Formation,WestFallsGroup)...... Second black band (of Clarke, 1898; Rhinestreet Shale)...... 43 ...... 1,4-5, 6, 49, 55-57, 58- SherburneSiltstone(Formation,GeneseeGroup)...... 59, 63, 67, 70, 79-80, 85, 184; Text-figs 2-3,4,16, 23-24, 63 ...... 4-5, 6, 12, 16, 21, 24-27, 99-100, 102, PipeCreekShale/HanoverShalecontact...... 5, 56; Text-figs 3, 16 210; Text-figs 2-3,4, 7, 28A; Pl.4,Figs7-9,Pl.5,Figs1-9 Platyceras Bed(WestBrookMember,TullyLimestone)...... ShurtleffSeptarianHorizon(SH;Upperdivision,CashaquaShale)...... 158, 165; Text-fig. 54K ..5-6, 35-38, 39-40, 41-42, 67-68, 79, 83, 103-105, 110, 125, PointBreezeGoniatiteBed(PB;AngolaShale)...... 127, 129-130, 157, 165, 177, 214, 217, 222, 225-226, 233-234, ...... 52-54, 55-57, 79, 84-86, 131, 258, 265; Text-figs 3-4, 10, 23, 30E-H, 31A, C-E, G-H, 32B, 134-144, 148, 166-169, 175-176, 179; Text-figs 15, 23, 41-44, 38C, 55C; Pl. 6, Figs 1-15, Pl. 7, Figs 4-6, Pl. 10, Figs 5-9, 11, 47, 53, 60; Pl. 16, Figs 3-8, Pl. 20, Figs 1-6, 9-10, Pl. 23, Figs Pl. 11, Figs 1-13, Pl. 12, Figs 8-10, Pl. 15, Figs 1-2, Pl. 16, Fig. 7-12, Pl. 32, Figs 4-9, 11-14, Pl. 33, Figs 3-4, 9-14, 19-20 11, Pl. 28, Figs 6-10, Pl. 31, Figs 18-20 Portage (Ithaca beds) at the University Quarry (Ithaca, Tompkins County) SonyeaGroup...... 4-5,6,8,17,34-35, 36-37, 49-50, ...... 108; Pl. 9, Fig. 1 63-64, 68-69, 79, 92-93, 95, 103-105, 108-110, 115, 129, 130, Portage(Naples)Beds...... 115, 225; Pl. 11, Figs 6, 12-13 132, 137, 157-158, 165, 177; Text-figs 2, 3-4, 10, 14, 18, 23 Portage (rocks, beds, shales, Group; outcrops in town of Naples; sandstone SonyeaGroup(majorrhythmABCBA)...... 69; Text-fig. 19A on East Hill, Naples, Ontario County; Java, Wyoming County; Sonyea Group equivalents (south and southeast of Ithaca, Tompkins Eighteenmile Creek, Erie County) ...... 3,34,62,95,123, County)...... 43,49-50; Text-fig. 14 131, 139, 153, 166; Text-figs 43-44, 50; Pl. 23, Figs 13-14 South Wales Shale (Canadaway Group) ...... 59 Portage Group (Ithaca, Tompkins County) ...... 108;Pl. 34, Fig. 5 SquawBayLimestone(Michigan)...... PortageGroup(Lower)(GeneseeRiver)...... 45,57 ...... 68,112-113, 117, 120, 158, 165-166, PortageSandstone(GeneseeRiver)(WestFallsGroup)...... 170, 254, 257; Text-fig. 54H-J; Pl. 26, Figs 8-9,Pl.27, Fig. 16 ...... 140, 148; Text-fig. 44 Starkeyblackshale(PennYanShale)...... 21 ProutLimestone(ErieCounty,Ohio)...... 81 Styliola Limestone(GenundewaLimestone)...... 27,115 HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 285

Table Rock Sandstone (Genesee River) (Gardeau Shale,West Falls Group) 34-35, 43, 51, 57, 65, 68-70, 79-80, 92, 95-98, 108, 123, 134- ...... 5-6, 45, 57, 148; Text-figs 3-4, 12 135, 137-142, 144-145, 148, 155, 161, 166, 178-179; Text-figs Taghanic Onlap (Sequence) . . . 5, 9, 12, 63, 66, 70, 186; Text-figs 3, 7 2-3, 4, 14, 23, 41A-G, 43A-g, 44A, C-F, 45B; Pl. 3, Fig. 4 Taghanic Unconformity ...... 9, 12-13; Text-fig. 7 West Falls Group (Formation), Lower (major rhythm ABCA) ...... Trinity Horizon (TR; “The Trinity” black shales; Angola Shale) ...... 69; Text-fig. 19B ...... 51-53, 55, 57, 68, 79, 85; Text-figs 15, 23 West Falls Group equivalents (south and southeast of Ithaca, Tompkins Tully Limestone (Formation) ...... County) ...... 49-50, 95, 97-98; Text-fig. 14; Pl. 3, Fig. 4 ...... 1-5, 6-7, 9-10, 11-12, 16, 24, 27, 63, 66, 70, 78-79, 80- West Hill Member/Nunda Sandstone interval (West Falls Group) ...... 82, 89-92, 158, 165, 170, 184, 186, 205; Text-figs 2-3, 4-5, 6- ...... 5-6, 45, 51, 55, 68; Text-figs 3-4 7, 23-24, 26D-H, 54K, P, 63; Pl. 1, Figs 1-8, Pl. 27, Fig. 11 West Hill Shale (Flags; Sandstone; Member, Angola Shale) ...... Tully Limestone (Upper Member/Division) ...... 5-6, 45, 51, 55, 68, 140, 148; Text-figs 3-4 ...... 9-10, 11-12, 27, 66, 70, 82, West River Shale (23 cycles/rhythms) . . . 30, 33, 69-70; Text-fig. 9 186, 205, 257; Text-figs 5-7; Pl. 1, Figs 1-8, Pl. 27, Fig. 11 West River Shale (Formation, Genesee Group) ...... 3-5, 6, 12, Tully Limestone disconformities (pre-intra-post Tully disconformities) . . 27, 30, 31-33, 35, 65, 67-69, 79-80, 82-83, 102-107, 113-115, ...... 9-10, 12, 78; Text-figs 5-7 117, 121-122, 129, 136, 164, 172, 184, 217-218, 270; Text-figs Tully Limestone-Geneseo Shale contact ...... 2-3, 4, 7, 9, 23-24, 30D, 33A-E, H, K-M, 34H, K, 58A-B, 63; ...... 5-6, 9-10, 12; Text-figs 3-4, 5-7 Pl. 7, Fig. 1, Pl. 8, Figs 10-11, 14, Pl. 34, Figs 1-4 Tully Pyrite (Leicester Pyrite) ...... 158-159, 163 Westfield Shale (Canadaway Group) ...... Union Spring Shale (Werneroceras Bed; Marcellus Formation, Hamilton ...... 4, 61, 180, 184, 262; Text-figs 2, 24; Pl. 30, Figs 1, 3 Group)...... 162 Widder Shale (Ontario, Canada) ...... 81-82, 254; Pl. 26, Fig. 5 Upper black shale (of Hall, 1843, of Reports; Rhinestreet Shale) . . 12, 43 Williams Brook Coquinite (Genesee Group) . . . . . 17, 24-25, 82, 108 Venango Sandstone, Lower (Cattaraugus; Conewango Group; Erie Williamsburgh Bed (Fossil Log Horizon, FL; West River Shale) ...... County, Pennsylvania) ...... 4, 62, 87; Text-fig. 2 ...... 31-32, 33, Virgin Hills Formation (Western Australia) ...... 150, 155 79, 102, 104-105, 122, 136; Text-figs 9, 23, 30D, 33K-L Volusia Member (Conneaut Group) ...... 62 Windom Shale (Member, Moscow Formation, Hamilton Group) ...... Wanakah Shale (Member, Ludlowville Formation, Hamilton Group) ...... 9,11- ...... 169 12, 17, 21, 158, 164, 257; Text-figs 7, 54D; Pl. 27, Figs 1-6 Werneroceras Bed (Union Springs Shale, Marcellus Formation, Hamilton Wiscoy Sandstone (and equivalents) (West Falls Group) ...... Group)...... 162 ...... 4-5, 6, 57-58, 59, 145, 148; Text-figs 2-3, 4, 45C-D West Brook Shale (Member, Bed, Tully Formation) ...... Wocklum (unit in Upper Devonian of Germany) ...... 77 ...... 9-10, 66, 70, 82, 91, 158, 165, Wolf Creek Conglomerate (Conewango Group) . . . . . 4, 62; Text-fig. 2 205, 257; Text-figs 5-6, 54K, P; Pl. 1, Figs 1-8, Pl. 27, Fig. 11 Woodmont Shale (or equivalents; West Virginia, Maryland) ...... 157 West Falls Group (Formation) ...... 4-5, 6, 8, 286 BULLETINS OF AMERICAN PALEONTOLOGY no. 374 HOUSE & KIRCHGASSER: LATE DEVONIAN GONIATITES 287 288 BULLETINS OF AMERICAN PALEONTOLOGY no. 374 PREPARATION OF MANUSCRIPTS

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