UNIVERSITY OF CINCINNATI
DATE: May 29, 2003
I, , Jocelyn Anne Sessa hereby submit this as part of the requirements for the degree of:
Master of Science in:
The Department of Geology It is entitled:
THE DYNAMICS OF RAPID, ASYNCHRONOUS BIOTIC
TURNOVER IN THE MIDDLE DEVONIAN APPALACHIAN
BASIN OF NEW YORK
Approved by:
Dr. Arnold I. Miller
Dr. Carlton E. Brett
Dr. David L. Meyer
Dr. Gordon C. Baird
THE DYNAMICS OF RAPID, ASYNCHRONOUS BIOTIC TURNOVER IN THE MIDDLE DEVONIAN APPALACHIAN BASIN OF NEW YORK
A thesis submitted to the
Division of Research and Advanced Studies Of the University of Cincinnati
in partial fulfillment of the requirements for the degree of
MASTER OF SCIENCE
in the Department of Geology of the College of Arts and Sciences
2003
by
Jocelyn Sessa
B.A. State University of New York at Geneseo, 2000
Committee Co-Chairs: Dr. Carlton E. Brett and Arnold I. Miller ABSTRACT
High resolution stratigraphic and paleoecological analyses of regional biotic turnovers can reveal much about their evolutionary and ecological processes. Biotic turnover in the
Devonian of the Appalachian Basin was investigated to determine whether taxonomic replacement across a key boundary was: 1) abrupt or gradual; 2) synchronous across the basin.
Analyses of abundance data that reveal a depth-controlled gradient persisted, with moderate variation, until just prior to the boundary. Two biotic assemblages occur above and below the boundary, while another occurs only above it. Turnover was asynchronous; certain assemblages appeared earlier than others. Above the boundary, a unique assemblage appeared abruptly, another assemblage appeared gradually, and a third remained unchanged.
Taxonomic composition was controlled by sedimentologic/environmental variables; specific faunas occur in particular lithofacies. Many taxa were not facies restricted, and occurred in many environments, but their abundance varied significantly. Taxon abundance and paleoenvironmental context is therefore important for understanding biotic transitions.
ii
iii ACKNOWLEDGEMENTS
I wish to sincerely thank my committee co-chairs, Dr. Arnie Miller and Dr. Carl Brett, for their
support and guidance throughout my studies at UC. Arnie, you have given me the tools to think
critically, pose interesting research questions, to balance work with everything else, and have
provided a role model I can aspire to. Thanks also for bringing me into the PBDB, and exposing
me to different ideas and research questions.
Carl, thank you for all of the fun hours in the field and looking at specimens, be it in the car
during thunderstorms or in the lab at night. Your unbridled enthusiasm and support of this
project is sincerely appreciated. Thanks also for introducing me to SSETI and giving me the opportunity to be part of such an interesting and fun experience.
Special thanks to my committee member Gordon Baird his hospitality on numerous occasions, for greatly assisting in fieldwork, and providing thoughtful comments on both research and music. Thanks to committee member Dave Meyer, whose thoughtful comments and general good nature are always appreciated. The Paleontological Research Institution in Ithaca New
York, and specifically Paul Harnik, were invaluable in collecting samples and providing assistance during fieldwork. Evelyn Mohalski Pence, graphic designer, was extremely helpful with the drafting of figures.
Fieldwork was aided greatly by the skills and time of Sue Taha-McLaughlin, Rich Krause, Alex
Bartholomew, and James Bonelli, who all provided companionship during sampling. A. Bart at the Center for the Study of Dynamic Cyclo-Stratigraphy of the Siluro-Devonian Appalachian
Basin of New York State provided unending hours of laughs and interesting discussion about the
Hamilton Group. To my fellow students, thank you for your friendship, patience, listening, and support throughout my masters. You will be missed.
iv To my parents and family, thank you all for your support and your encouragement to pursue my dreams.
To James, I could not have made it through without you. Thanks for your friendship, compassion, and love – you are my sunshine.
Funding for this research was granted through Sigma Xi, the American Museum of Natural
History Theodore Roosevelt Fund, the Paleontological Society, the Geological Society of
America, a University of Cincinnati Student Summer Research Grant, a University of Cincinnati
Office of Sponsored Research matching grant for Sigma Xi awards, a University of Cincinnati
Departmental Fellowship, and NASA Exobiology Grants awarded to Dr. A.I. Miller.
v TABLE OF CONTENTS
Page
ABSTRACT…………………………………………………………………………..….ii
ACKNOWLEDGEMENTS………………………………………………………………iv
LIST OF FIGURES.………………………………………………………………………2
LIST OF TABLES………………………………………………………………………...3
LIST OF APPENDICES…………………………………………………………………..4
INTRODUCTION…………………………..…………………………………………….5
GEOLOGIC SETTING AND STRATIGRAPHY…...... …………………………………9
METHODS…..…..………………………………………………………………………19
RESULTS AND DISCUSSION…………………………………………………………23 Windom Samples…………………………………………………………….…23 Transitional Samples……………………………………………………..….…30 Tully Samples………………………………………………………………..…38 Summary of Results…………………………………………………………....46 Implications………………………………………………………………….…47
CONCLUSIONS…………………………………………………………………………52
REFERENCES…………………………………………………………………………...54
APPENDIX I……………………………………………………………………………..63
APPENDIX II………………….…………………………………………………………67
1 LIST OF FIGURES
Figure Page
1. Map of the study interval in New York State...... ……………………………….10
2. Schematic cross section of the eastern Appalachian Basin during uppermost Windom and Tully time………………………………………………………….11
3. General stratigraphic column of the study interval………………………………14
4. Detrended Correspondence Analysis of Windom samples and taxa…………….24
5. Abundance and occurrence graphs of key Windom taxa………………………..25
6. Abundance and occurrence graphs of key Highland Forest taxa………………..31
7. Detrended Correspondence Analysis Axis one scores plotted in stratigraphic order for Locality 10…..………..………………………………………………………35
8. Detrended Correspondence Analysis Axis one 5 point moving average scores plotted in stratigraphic order for Locality 10…………….………………………37
9. Detrended Correspondence Analysis for all samples and taxa…….…………….39
10. Abundance and occurrence graphs of key Tully taxa……………………………40
11. Summary figure with samples coded to Detrended Correspondence Analysis group …………………………………………………………………..41
12. Percent occurrence of the most common taxa within the studied interval………42
2 LIST OF TABLES
Table Page
1. Percentage of taxa within systems tracts of the Windom……………………….29
2. Percentage of key taxa from the Emanuella-dominated assemblage that occur in non-boundary Hamilton samples……...…………………….…….…...33
3 LIST OF APPENDICES
Appendix Page
I. Locality register and sample positions………………………….……………….63
II. Middle Devonian uppermost Hamilton Group and Tully Formation field census raw data…………………………………………………………...…...... 67
4 INTRODUCTION
Previous studies of ancient biotic transitions at regional scales have typically documented prolonged periods of relative compositional stability interspersed with relatively brief intervals of rapid biotic reorganization. In this context, many investigators have focused on delineating and calibrating the periods of stability (i.e., Boucot, 1983; Brett and Baird, 1995; Bennington and
Bambach, 1996; Holterhoff, 1996; Pandolfi, 1996; Sheehan, 1996; Patzkowsky and Holland,
1997; Olszewski and Patzkowsky, 2001; Gardiner, 2001). However, comparatively less attention has focused on the equally important intervals of biotic change (see Ivany, 1996, 1999, for theoretical discussions of turnover and its implications). High resolution analyses of critical boundaries can lead to a greater understanding of how, and perhaps why, these transitions occur.
Specifically, transitional patterns may be important in the diagnosis of interaction – or lack thereof - among species that compose assemblages, and the constraints placed upon organisms by their environments in response to physical changes. A detailed study of turnover events may also reveal the variables, be they biotic or abiotic, to which organisms respond most significantly on a regional scale.
Causal mechanisms aside, it would also be useful to identify just how abrupt turnover intervals are. Do turnovers take place across single horizons, or are they more gradual, with a progressive breakdown of existing assemblages and piecemeal development of subsequent biotas? It is plausible that these different end members could relate to the degree of biotic interaction within ‘communities’. If species within communities are strongly tied to one another by biotic interactions (e.g. Elton, 1927), then turnover should appear abrupt, given the unitary nature of the community. Alternatively, if species are not tightly linked to one another (e.g.
Gleason, 1926), then a more diffuse transition might be observed. Admittedly, these are not the
5 only possibilities, as abrupt turnover may simply be related to the shared environmental preferences of taxa that appear or disappear during a turnover event (sensu Miller, 1997). Thus, an understanding of each taxon’s environmental preferences would be helpful in teasing apart the processes responsible for biotic turnover, and necessitates the characterization of species abundance within various biofacies before, during, and after a turnover event.
It is also important to determine whether turnover is synchronous across coeval environments within a given region. Faunas in dissimilar environments are often thought to respond differently to a given disturbance (e.g., see discussions by Bretsky and Lorenz, 1970, and Allen and Sanders, 1996), and, thus, coeval assemblages in different facies may exhibit change to differing extents or at different times. If asynchroniety is found, detailed comparisons among biofacies may point to the causes of the turnover, and may help explain why one biota changed at a rate different than another. It should be noted that this is independent of whether faunal turnover was abrupt or gradual, and that several different permutations of these hypotheses are possible (e.g., abrupt and synchronous, abrupt and asynchronous, etc.).
To better understand the nature of regional biotic turnover, the present study focuses in detail on faunal changes within a key interval in the upper Middle Devonian Appalachian Basin of New York State. Specifically, transitions from biofacies of the uppermost Windom Shale of the Moscow Formation of the Hamilton Group to those of the overlying Tully Formation were evaluated in detail. This middle Givetian interval preserves a critical biotic transition both locally and regionally. Locally, many common and characteristic Hamilton Group taxa exhibit an apparent hiatus at the basal Tully as they migrated out of the Appalachian Basin, while previously unrecorded Tully taxa immigrated into it. This boundary was originally recognized worldwide as the beginning of both the ‘Taghanic Onlap’ (Johnson, 1970) a major sea level rise,
6 and later as the ‘Taghanic Event’ (House, 1985), a multiphased succession of extinctions that, in
total, may have been at least as severe as the Frasnian/Fammenian extinction (Aboussalam and
Becker, 2001; House, 2001). This bioevent was subdivided within the Appalachian Basin into
the Lower, Middle, and Upper 'Tully Bioevents' (Baird and Brett, 2003).
This study focuses primarily on the 'Lower Tully Bioevent' (Baird and Brett, 2003),
which begins at the basal Tully, during a time when sea-level highstand is associated with
basinwide changes in sediment composition and an apparent incursion of an “exotic” fauna from
the Old World Realm. Thus, the Hamilton/Tully transition provides an ideal boundary to observe
regional turnover dynamics. Until recently, a detailed microstratigraphic study of this transition
was not possible because both the uppermost Hamilton and lowermost Tully are absent due to an
unconformity in many areas. However, recent re-examination of several outcrops by Brett and
Baird (2001) revealed a nearly complete section of this critical interval and provided the focus for this study. These outcrops appear to record at least two slightly different facies/environments with distinct assemblages immediately before and after the boundary, allowing for tests of synchroniety among biofacies.
The lithology, sequence stratigraphy, and faunas of these units have been studied extensively (Hall, 1843; Grabau, 1917; Cooper, 1930; Grasso, 1973, 1986; Baird and Brett,
1983; Miller, 1986; Brower and Nye, 1991; Lafferty et al., 1994; Mayer et al., 1994; Brett and
Baird, 1994, 1996, 1997; Bezusko, 2001). This preexisting framework allows for the designation
of biofacies membership and general environmental preferences of taxa. Additionally, multiple
paleo-environments are available for study within the Windom and Tully, which permits
diagnosis of the environmental preferences of taxa both before and after the Lower Tully
Bioevent.
7 This study has two specific goals: first, to determine whether turnover from Windom biofacies to those of the Tully was abrupt, or whether Windom faunal elements gradually became less abundant while those of the Tully became more dominant; and second, to determine whether turnover occurred synchronously across the various facies present in this portion of the
Appalachian Basin at the end of Windom deposition. Further, the occurrences and peak abundances of various taxa are assessed to better understand the variables that contributed to faunal turnover.
8 GEOLOGIC SETTING AND STRATIGRAPHY
This study focuses on faunal changes near the stratigraphic and lithologic boundary
between the Hamilton Group and the Tully Formation, and the strata several meters below and
above this boundary in the Appalachian Basin of New York State (Figs. 1 & 2). This boundary
has worldwide significance, as it marks the beginning of global environmental change and
protracted, stepwise faunal extinction (House, 1971; Hallam and Wignall, 1997; Aboussalam and
Becker, 2000). Sea level rise begins at this boundary and has been associated with extinctions of many Paleozoic taxa, including brachiopods, corals, and ammonoids (House, 1985; Cooper,
1986). Sea level events have been termed the 'Taghanic Onlap' by Johnson (1970) and the associated biotic changes termed 'Taghanic Event' by House (1985). This extinction has not been investigated as much as other Devonian extinctions (e.g. the Late Devonian
Frasnian/Fammenian extinction) even though it may have reached 'mass' extinction levels
(Hallam and Wignall, 1997; Aboussalam and Becker, 2000, 2001; House 2001).
Further work has shown that, within the Appalachian Basin, the Taghanic Event can be subdivided into a series of events. The 'Lower Tully Bioevent' (Baird and Brett, 2003) commences with the deposition of the basal Tully during highstand conditions that may have been associated with dysoxia (Baird and Brett, 2003). The middle and upper events are subsequent faunal turnovers, with the upper event marking the final demise of many Hamilton and Tully taxa. The Lower Tully Bioevent has been tentatively recognized worldwide, although precise correlations have proven difficult (Aboussalam and Becker, 2000, 2001).
During the studied interval, uplift associated with the Acadian Orogeny produced siliciclastic sediments that were shed into the Appalachian Foreland Basin at a time when New
9
10
11 York State was situated about 25 to 30 degrees south of the paleo-equator. During the Late
Givetian (about 380 ma, Middle Polygnathus varcus conodont zone) sediments deposited on the
western flank of the basin (westward of the present day Finger Lakes) were comprised primarily
of calcareous shales and mudstones. Equivalent strata to the east, near Syracuse, are consistently
thicker and are composed of silty mudstones and fine grained sandstones. In much of western
New York, parts of the uppermost Windom and lower Tully are absent because of submarine
erosion and/or extended periods of nondeposition (Baird, 1981; Baird and Brett, 1981). This study focuses on the silty mudstone and mixed siliciclastic-carbonate facies of the northeastern portion of the Appalachian Basin near Syracuse, where sedimentation was fairly constant
through the entire interval represented in Figure 2.
The paleoshoreline of the Appalachian Basin trended roughly northwest to southeast, and
typically central New York was deeper than the eastern region. Overprinted on this general
eastward shallowing is a localized area of subsidence and deepening (Heckel, 1973), recently
termed the 'New Lisbon sub-basin' (Baird and Brett, 2003), that affected the lower Tully
Formation in the study area (Fig. 2). Thus, in the study area, deeper water environments of the
Tully are located to the east and slightly shallower environments lie westward. Sampled
localities run roughly parallel to the paleoslope of both the New Lisbon sub-basin and the
Appalachian Basin and thus, follow hypothesized onshore to offshore transects.
Absolute age dates for the Hamilton Group and Tully Formation are poorly known (Brett and Baird, 1996). Using cyclostratigraphy, House (1995) concluded that the entire middle varcus zone, of which this study encompasses only part, was about 0.9 Ma in duration. Additionally, given that about four, fourth-order cycles were studied (Brett and Baird, 1996) and each fourth order cycle represents 100 to 400 thousand years (Van Wagoner et al., 1990), the interval from
12 the upper Hamilton through the Lower Tully probably represents about 0.5 million years in
duration.
Sequence stratigraphic interpretation within the Middle Devonian is facilitated by numerous well exposed outcrops and simple geologic structure within New York State. Brett and Baird (1985, 1986, 1996) have used the sequence stratigraphic paradigm proposed by Vail et al. (1977, 1991) and Van Wagoner et al. (1988, 1990) to delineate different orders of cyclicity within the Hamilton Group and, more recently, the Tully Formation (Baird and Brett, 2003).
Cycles have been identified using various sources of information, including Waltherian relationships, sediment grain size and composition changes, and correlation of regional event beds. Large scale, third order sequences (0.5-3 ma; Vail et al., 1991) correlate approximately to formations (e.g. Moscow and Tully Formations), and are bounded by thin, widespread transgressive limestones at their bases and unconformities at their tops (recall that the Hamilton to Tully transition is variably disconformable throughout New York except in the study area,
Baird and Brett, 2003). Cycles are observed to vary predictably in thickness, cycle symmetry, and facies content across the basin topography, with thinner, more symmetrical cycles in the west (near Buffalo) and thicker, asymmetrical cycles in the east (south of Utica) (Brett and
Baird, 1985, 1986, 1996). Descriptions herein focus on the siltier, sub-symmetric cycles of the study area.
Within the third order Moscow Formation, small-scale fourth through sixth order cycles have been identified. These cycles have been designated informally with bed names by Brett and
Baird (1994, 1996); all Windom Shale bed names in text and figures are informal. This study focuses on the fourth order cycles, also called parasequence sets or subsequences. Specifically, part of a lower subsequence, two succeeding complete subsequences, and an uppermost,
13
Figure 3.Generalstratigraphic columnofthestudyinterval,withsequencestratigraphic interpretations. Hamilton Group Tully Formation Moscow Formation Lower Mid. Upper
Bear SwampBed Amsdell Bed Fisher GullyBed Fall BrookBed T South LansingBed Spezzano Bed Gage GullyBed Sheds Bed Highland ForestBed DeRuyter Bed Cuyler Bed Fabius Bed T Carpenters FallsBed T W Karla Bed Karla aunton Bed aughannock FallsBed ully V est BrookShaleBed alley Bed Scale (m) 10 12 14 16 18 20 22 24 26 28 6 8 0 2 4 HST? TST? MFS MFS HST RST HST RST RT TST TST S Cycle 1 Cycle 2 Mudstone Siltstone Limestone Silty Mudstone Concretions Fossiliferous LS Unconformity KEY 14 incomplete subsequence of the upper Windom and one subsequence of the Tully were analyzed
(Figs. 2 and 3).
In the study area, the Windom Shale consists primarily of gray, fossiliferous mudstones, and records a range of environments, from shallow, sandy shelves to deeper basinal settings.
Widespread, thin calcareous mudstone shell beds that are locally coral rich (e.g. lowest Amsdell and South Lansing beds) are easily recognized and correlated within Windom strata, and form the bounding units of fourth order cycles, as inferred by sequence stratigraphic study (Brett and
Baird, 1994, 1996). These shell beds are interpreted to represent the beginnings of transgressive systems tracts (TSTs), and are the shallowest Windom facies observed (Brett and Baird, 1985).
Calcareous shales of the TST lie above these beds (e.g. middle Amsdell and Spezzano beds), and are interpreted to represent oxygenated, low turbidity environments (Brett and Baird, 1985;
Vogel et al., 1987; Linsley, 1991).
TST deposits are overlain by maximum flooding surfaces (MFSs), which represent the deepest water conditions found within the Windom. These beds (e.g. upper Amsdell and Karla beds) record periods of minor sediment starvation and are persistent and roughly isochronous across the basin (Brett and Baird, 1986, 1996). This, combined with the ease of recognition in outcrop, has made MFSs extremely useful for regional correlation (Brett and Baird, 1986, 1996).
Early highstand deposits (e.g. Fisher Gully and Gage Gully beds), lie above the MFSs, and are represented by dark gray, fissile to laminated chippy shales that contain pyrite and are interpreted to have been deposited near the dysoxic-anoxic boundary (Grasso, 1973; Brower et al., 1978; Kammer et al., 1986; Thompson and Newton, 1987; Vogel et al., 1987; Brett et. al,
1990; Brower and Nye, 1991).
15 These shales are typically succeeded by gray, Zoophycos-burrowed mudrocks and siltstones of the late highstand or regressive systems tract (RST). RST deposits (e.g. Bear
Swamp, Taunton, and Sheds beds) are interpreted to have been deposited at intermediate depths above storm wave base on an otherwise low energy ramp (Grasso, 1973; Brett et. al, 1990;
Linsley et al., 1992; Newman et al., 1992). Gray silty mudstones comprise the bulk of the
Windom Shale and typically underlie TST shell beds.
Within most of New York State, the Gage Gully bed is typically unconformably overlain by the Tully Formation. Cooper and Williams (1935) and Heckel (1973), recognized the diachronous nature of the Hamilton/Tully contact, but also described localities near Syracuse and eastward where the boundary is more conformable. Brett and Baird (2001) have studied this area in detail (highlighted in Fig. 1), and have identified portions of an additional fourth order sequence, as well as informally naming two units that overlie the Gage Gully bed. The lower of these is the Sheds bed, a probable RST deposit that locally contains abundant fossils and is lithologically similar to the Taunton bed. The Sheds is overlain unconformably by the Tully in some localities. The Highland Forest bed (so named for Locality 10 on Fig. 1, Highland Forest
Park, where it reaches maximum thickness) is above the Sheds, and, at some localities, this transition is marked by a shell lag, which may represent the transition from RST to TST. At some outcrops the Highland Forest contains minor, discrete siltstone beds near the top. Pyrite burrows and phosphate nodules occur throughout the unit (personal observation). During the end of Highland Forest deposition, siliciclastic sedimentation waned, while carbonate production increased. This change to a mixed siliciclastic and carbonate system makes precise sequence stratigraphic interpretations difficult, but the Highland Forest is thought to represent TST and
16 HST conditions. Additionally, it appears that the Highland Forest to Tully contact is probably
slightly disconformable (Baird and Brett, 2003; Bartholomew, 2002).
In the study area, the Tully Formation is composed primarily of muddy siltstones and
quartz sandstones containing variable amounts of carbonate (Heckel, 1973). It was deposited
during a period of tectonic quiescence from the Acadian Orogeny (Ettensohn, 1985), and is
composed of sediments deposited primarily in deep shelf environments (Baird and Brett, 2003).
The Tully appears to represent a third order sequence that contains three fourth order sequences respectively named the Lower, Middle and Upper Tully (Baird and Brett, 2003).
In the present study, the basal unit of the Tully (DeRuyter Bed), was sampled extensively, as it directly overlies the Windom. It is a fossiliferous, calcareous siltstone that contains pyrite and sporadic phosphate nodules, and chamosite “ooids” in eastern localities that are visible in outcrop (Heckel, 1973). These minerals are interpreted to have formed during periods of sediment starvation, resulting from a minor transgression and highstand (Baird and
Brett, 2003). Most upper Lower Tully units are commonly eroded at a mid-Tully unconformity in the study area, and only a few samples from upper Lower Tully units were included in statistical analyses.
Middle Tully units rarely contain fossils in the study area, and were not included in statistical analyses. The Middle Tully is overlain by the 50 cm thick West Brook Shale, a horizon rich in fossils, which comprises the top of the studied interval.
It should be emphasized that the Hamilton to Tully faunal transition was ephemeral in some respects, as many common Hamilton taxa recur in the overlying West Brook Shale Bed
(Cooper and Williams, 1935; Bonelli et al., 2003). Four samples from the West Brook Shale were included in one analysis, but were not the focus of the present study.
17 The fauna found within Hamilton strata of New York have been studied extensively in a
paleoecologic context, both qualitatively and semi-quantitatively (Baird and Brett, 1983; Brett et.
al, 1990; Brett and Baird, 1995) and with quantitative methods (e.g. NonMetric
MultiDimensional Scaling by Brower et al., 1978; Brower, 1987; Brower and Nye, 1991;
Correspondence Analysis by Newman et al., 1992; polar ordination by Lafferty et al, 1994; and
Detrended Correspondence Analysis by McCollum, 1991; Bezusko, 2001). In general, biotas
have been shown to vary in direct association with lithologic changes. Faunas are thought to
have been controlled by bathymetric gradients related to sedimentation, decreasing oxygen
content, and storm frequency (Savarese et al., 1986; Brett et. al, 1990; Brower and Nye, 1991).
Faunal density of most Lower Tully units is markedly less than that of the underlying
Windom. This, combined with the hard splintery lithologic nature of the Tully, has made prior faunal investigations difficult (but see Heckel, 1973, for general faunal characteristics).
18 METHODS
Samples were taken from the 21 localities illustrated in Figure 1. Individual counts of
organisms, either from bedding planes or a roughly 1,000 cubic cm interval of rock, were taken
in the field from the strata shown in Figures 2 and 3. Special care was taken to sample all
fossiliferous bedding planes. Samples were spaced roughly half a meter to one meter apart
stratigraphically where fossil abundance and distribution permitted, as certain units are more
fossiliferous than others. Spacing between samples near the Windom/Tully boundary was
typically a maximum of 10 cm apart.
One locality (Locality 10 on Fig.1, Highland Forest Park) was found to contain the most
complete transition between the Hamilton and Tully, and therefore was the most intensely
sampled locality in this study. Sampling here was aided by volunteers from the Paleontological
Research Institution, and samples were used in a related project (Harnik and Ross, 2003).
It is particularly important to quantify paleoecologic changes in terms of abundance, as many species are widespread among samples but vary significantly in abundance. Most species display Gaussian distributions (Whittaker, 1975; Holland 1995, 1996), and several ecologic and paleoecologic studies have recognized that the use of abundance versus presence/absence data will affect the designation of communities (Jackson 1994; Pandolfi, 1996; Olszewski and
Patzkowsky, 2001). Statistically recurring assemblages have been found to be more difficult to document using abundance data (Rahel, 1990). In this study abundance data were used to adequately characterize environmental tolerances and preferred habitats of taxa, and to provide a more stringent test of recurrence.
19 To help ensure sample standardization, a collection curve of taxa versus specimens was
generated for each sample (see Colwell and Coddington, 1994; Gotelli and Colwell, 2001, for a
theoretical discussion of this and other sampling techniques). Individuals were counted until the
collection curve ‘leveled off’ i.e., when 10 successive individuals were counted without
encountering any new taxa.
For brachiopods and bivalves (83% of all taxa) one disarticulated valve or two articulated
valves were counted as one individual. Bryozoans and crinoid columnals were categorized as
rare, common, or abundant, which were later assigned values of one, three, and six, respectively, for multivariate statistical analyses, given that these classes approximately represent logarithmic differences in abundance (see Miller et al., 2001; Holland et al., 2001; Webber, 2002a, for a discussion of a similar approximation).
Brachiopod specimens were identified to the genus level in nearly all cases and to species level where possible. As is common in paleoecologic studies, analyses were conducted using genus level identifications, to incorporate the majority of data and to effectively capture paleoecologic signals. This distinction is immaterial in any case, as most brachiopod genera within the Appalachian Basin appear to have been monospecific through the upper Middle
Devonian. Family level categories were also utilized for fragmented or poorly preserved specimens; combined these categories comprise 3% of all individuals. Analyses were run including and excluding family data, and results were robust to these treatments. The low abundance of these categories indicates minimal taphonomic bias, via fragmentation or reworking, present in the study.
Bivalves were identified to species level where possible. Bivalve genera are also largely monospecific within the study interval, with the exception of the common genus Paleoneilo,
20 which has three species that are easily distinguished. Thus, for analyses, species of Paleoneilo were used, rather than the genus. Gastropods were rare within the study interval and were identified to genus level when possible. Fragmentary and unidentifiable specimens were placed in the ‘gastropod fragments’ category, which comprised less than 1% of sampled specimens.
Crinoids were found only as disarticulated, scattered columnals, and, as such, were placed into an “unidentifiable crinoid” category. Complete trilobites were identified to genus level, while isolated cephalon, thorax, or pygdia were placed into an “unidentifiable/fragmented trilobite” category. Corals were identified only coarsely; i.e. rugose, tabulate, or auloporid. Bryozoans were identified either to family or morphotype. Taxonomic identification, particularly for brachiopods and bivalves, was based primarily on Linsley, 1994.
Taxon abundances for each sample in the data matrix were transformed to percentages to help mitigate differences in the total number of specimens from sample to sample. This matrix was then analyzed using a variety of multivariate statistical techniques, including cluster analysis with Unweighted Pair Group Method with Arithmetic Averaging (UPGMA) (Sneath and Sokal,
1973), Detrended Correspondence Analysis (DCA) (Hill and Gauch, 1980), and Nonmetric
MultiDimensional Scaling (NMS) (Kruskal, 1964; Clarke, 1993). All analyses used the Bray-
Curtis similarity coefficient (Bray and Curtis, 1957), and were conducted using PC-ORD4
(McCure and Mefford, 1999).
Subsidiary data for samples, including lithology, stratigraphic position, and location within parasequence set, were used to interpret outcomes of multivariate analyses. Inferred trophic role (e.g. burrower, suspension feeder) described in Thayer, 1975, and higher order taxonomic membership of taxa were also utilized for this purpose.
21 Only DCA results are presented in this paper, as cluster analysis and NMS results are in agreement with those of DCA. DCA was designed by ecologists to analyze faunal compositional changes along well constrained physical gradients (Hill and Gauch, 1980). Several marine paleoecologic studies have shown that the ordination of samples along Axis 1 tends to be associated with environmental variables correlated with water depth, such as turbidity, light level, or frequency of storm disturbance (Patzkowsky, 1995; Holland et al., 2001; Miller et al.
2001; Bezusko, 2001; Webber, 2002a). As shown below, DCA was a useful tool for understanding the faunal relationships present within and among the various biofacies present in the Windom and Tully.
DCA compares samples to one another by taxonomic composition and (in this study) abundance, while simultaneously comparing taxa to one another by their presence and abundance in samples. Samples and taxa are arrayed along axes that illustrate their relative similarities and differences. Therefore, taxa and samples that occur in the same area of an ordination are similar to one another. Although both samples and taxa scores can be plotted on one graph, for simplicity, in the presentation below “A” figures display sample scores for axes 1 and 2 and “B” figures show taxon scores for the same axes.
22 RESULTS AND DISCUSSION
Windom Samples
A 152 sample by 111 taxon data matrix composed of about 4,900 individuals was
analyzed in this study (Appendix II). Sample size ranged from 11 to 77 specimens, with an
average of 31. While the size of the least numerous samples may seem low, it appears to
accurately characterize the taxonomic composition of some beds, as discussed below.
Multiple DCA trials were conducted on variously culled data sets that excluded certain
categories and/or taxa from the data matrix to assess the response of sample and taxonomic
patterns. In one trial, taxa were culled so that only the 25 most abundant remained in the data
matrix. Sample ordinations for this analysis were virtually identical to those found using the
entire data matrix. Analyses were also performed excluding all fragmentary or unknown categories (including the “unidentifiable crinoid” category) and again, patterns remained virtually unchanged. These permutations indicate the robustness of the patterns contained within the data matrix.
Before comparing faunas of the Windom and Tully, or focusing on the transition interval between the two, it is useful to first concentrate on the Windom Shale, to understand faunal dynamics during the “background” interval immediately prior to the transition. For this purpose, a DCA of Windom data only is presented in Figure 4. In Figure 4A samples were coded according to their position within systems tracts. Key Windom taxa are highlighted in Figure
4B. Samples from the uppermost Windom Highland Forest bed were included in this analysis, and are highlighted in Figure 4A, as they are of particular interest, and will be discussed shortly.
Abundance graphs for key taxa among systems tracts are given in Figure 5. Figure 5B displays
23 DCA of Windom Samples 500 A TST RST HST and MFS 400 Highland Forest low dominance assemblage Highland Forest Emanuella- 300 dominated assemblage
200
100
0
Axis 2 DCA of Windom Taxa 500 B
400 Emanuella
Mucrospirifer
300
Fenestrate Bryozoa Megastrophia Modiomorpha Eumetabolotoecia Indet Crinoids 200 Tropidoleptus Phestia Pseudoatrypa
Nuculoidea Rugosa Longispina
Actinopteria Mesoleptostrophia Devonochonetes P. emarginata 100 P. constricta
Nuculites P. filosa
Allanella 0 -100 0 100 200 300 400 500 Axis 1 Figure 4. DCA axis one and two scores for Windom samples (A) and taxa (B). Samples have been coded according to position in parasequence cycles. The uppermost Windom samples are distinguished from RST and TST deposits in A. Several key taxa are highlighted in B.
24 ABHST Samples (n = 46) Megastrophia RST Samples (n = 28) Mesoleptostrophia TST Samples (n = 24) Corals
Bryozoa
Tropidoleptus
Mediospirifer
Indet Crinoid
Rhipidomella
Pustulatia
P. emarginata
Longispina
Mucrospirifer
P. constricta
Devonochonetes
Nuculoidea
Emanuella
Allanella
Eumetabolatoechia
0 1020304050 10 20 30 40 50 60 70 80 90 100 Percent Abundance Percent Occurrence Figure 5. Graphs of key Windom taxa showing A) Each taxon's average percent abundance when it occurs in each systems track and B) Percentage of samples within each system track that a taxon occurs in. Corals = Rugosa + Tabulate + Auloporid, Bryozoa = Fenestrate + Sulcoretepora.
25 the percentage of samples within a given systems tract that a taxon occurs in, and Figure 5A
gives the average abundance of a taxon when it occurs in that systems tract (not a taxon's abundance throughout all samples of a systems tract).
Collectively, Windom samples are arrayed continuously along Axis 1, with no breaks between sample groups. This pattern implies a faunal gradient along Axis 1, produced by taxa with broadly overlapping ranges throughout samples. This axis correlates very well to sample position within cycles, and thus may reflect environmental changes related to sedimentary cycles and bathymetric change. Samples with low scores (i.e., to the left) on Axis 1 are from dark gray, chippy, pyritic shales of the upper Amsdell, Fisher Gully, Gage Gully, and Karla beds that comprise HST and MFS deposits. Figures 4B and 5 indicate that these deposits are
overwhelmingly dominated by the leiorhynchid brachiopod Eumetabolotoechia multicostum and
the small spiriferid brachiopod Allanella sp.; in the field, monospecific pavements of
Eumetabolotoechia are very common. In fact, each brachiopod was found to comprise 100% of a few samples in this biofacies, producing a separation along Axis 2 of samples on the left side of
Axis 1 (Fig. 4A). These samples contain the smallest number of individuals (11) within the study. This sample size faithfully represents the monospecific nature of these deposits, as additional specimens were counted in the field and no new taxa were encountered; only 11 individuals were reported so that the sampling procedure described in the methods section remained consistent.
Many studies of Devonian biofacies in general, and of Hamilton biofacies in particular, have found Eumetabolotoechia in association with dark gray shales deposited under dysoxic conditions (Kammer et al., 1986; Brower 1987; Thompson and Newton, 1987; Brett et al, 1990;
Brett et al. 1991; Brower and Nye, 1991; Gray, 1991; Linsley, 1991; McCollum, 1991; Newman
26 et al., 1992). Interestingly, the specimens of Allanella sp. are diminutive as compared to
Allanella tullia, which is commonly found in shallow water sandstones of the Hamilton (Brett et al., 1990). It is possible that Allanella sp. may represent a juvenile form of Allanella tullia
(Harnik et al., 2002), although it is unclear why small specimens have been found only within the dysoxic facies of these upper Hamilton deposits.
The next set of samples encountered along the gradient are medium gray, Zoophycos bioturbated mudstones and siltstones of the Bear Swamp, Taunton, and Sheds beds, interpreted to represent shallowing or regressive portions of cycles, and the Fall Brook bed, interpreted as the transition from HST to RST conditions. These samples are of intermediate diversity, unlike the HST samples, and are not as strongly dominated by just one or two taxa (Fig. 5). Of all biofacies studied, this has the highest percentage of bivalves, especially infaunal forms, although no particular genus becomes abundant. Common infaunal bivalves highlighted in Figure 4B are
Paleoneilo constricta, P. filosa, P. emarginata, Nuculites, and Nuculoidea. The DCA values of these nuculid bivalves are transitional between the HST and RST deposits, according well with the recognition that nuculids can tolerate upper dysoxic conditions (Dick and Brett, 1986;
Kammer et al., 1986, Brett, 1998). Examples of additional bivalves consistently present in low abundance include Modiomorpha, Actinopteria, and Phestia. Notice that these bivalves plot farther to the right along Axis 1 (Fig. 4B), indicating they are not present within HST deposits, and were probably unable to tolerate dysoxic conditions.
Brett et al. (1991) compared highstand faunas of dark gray to black, fissile shale deposits to those from regressive, grey bioturbated mudstones within the Windom Shale. They found faunal patterns similar to this study: a greater diversity of bivalves in the grey mudstone RST facies, and higher dominance with reduced overall diversity in the dark gray shales of the HST.
27 They attributed these differences to the dysoxic conditions of the dark gray shales versus the
more oxygentated conditions suggested by the bioturbated mudstones.
Samples that plot to the far right on Axis 1 are composed of light gray mudstones, shelly limestones and local coral beds of the lower Amsdell, Lansing, and Spezzano beds, interpreted as the initiations of transgressions, as well as several Taunton and Sheds samples, which are associated with RSTs. These samples are characterized by high diversity, low dominance brachiopod rich assemblages typical of shallow, oxygenated environments. The brachiopods
Tropidoleptus and Mediospirifer are the two most abundant taxa found in this biofacies, comprising about 30% of the total genus composition (Fig. 5). Crinoids, corals, and bryozoans are also common, and have their highest abundances in these TST samples.
Gray (1991) studied a shallowing-up cycle within the middle Hamilton Group (the
Centerfield Member) and recognized patterns similar to those diagnosed here. The cycle he analyzed begins with dysoxic, Eumetabolotoechia-dominated shales and culminates with a diverse brachiopod assemblage characterized by Mediospirifer, Tropidoleptus, and corals. Vogel et al. (1987) studied faunal assemblages throughout the Hamilton Group and also reported similar biofacies associations, from ‘Leiorhynchus’ (now Eumetabolotoechia) dominated black to dark gray shales, to a Mucrospirifer – chonetid – bivalve-assemblage in medium gray silty mudstones, and a shallow water, high diversity biofacies, found in light gray calcareous mudstones.
Focusing more specifically on taxonomic distributions among systems tracts, an interesting pattern is observed when comparing the abundances of taxa that occur in multiple systems tracts versus those that are restricted to a single system tract. Table 1 indicates the percentages of taxa found in one, two, or three systems tracts. Among all taxa, 35% occur in all
28 All 3 STs 35% 2 STs 26% 1 ST 38% TST & RST 15% TST Only 14% TST & HST 3% RST Only 17% RST & HST 8% HST Only 7%
Table 1. Percentage of Hamilton taxa that occur in various systems tracks (STs).
29 three systems tracts, and thus, in several different facies. If we consider just the key taxa illustrated in Figure 5, this number increases to 78% (14 out of 18). This reinforces the suggestion that most of the studied taxa had broad environmental ranges and tolerated a variety of conditions, and highlights the importance of relative abundance data for diagnosing biotic variations within and among facies. Several other quantitative studies of Hamilton biotas reported similarly large ranges for common taxa (Brower et al., 1978; Savarese et al., 1986;
Brower & Nye, 1991; McCollum, 1991; Newman et al., 1992; Lafferty et al., 1994).
Transgressive deposits contain a high number of taxa unique to a single systems tract, including the brachiopods Megastrophia and Mesoleptostrophia (Figs. 4B and 5). These taxa have been noted as facies restricted in other Middle Devonian studies (Brett et al., 1990) and a concentration of first occurrences of shallow water taxa is expected at basal TSTs based on theoretical modeling, because of nondeposition at sequence boundaries and because of large facies changes at the bases of TSTs (Holland, 1995, 1996).
Transitional Samples
Returning to Figure 4A, two distinct groups of uppermost Windom (Highland Forest bed) samples are seen - those that plot within the Windom gradient (called ‘low dominance Highland
Forest’ in Fig. 4A) and those that group outside of it (called ‘Emanuella-dominated Highland
Forest’ in Fig. 4A). Figure 6 shows the most abundant taxa for each of the Highland Forest groups. Low dominance Highland Forest samples are compositionally similar to previous RST assemblages described, and can be thought of as a ‘typical’ Windom assemblage. The
Emaunella-dominated samples also contain taxa that were found to occur throughout the
Windom; only one taxon in this group is not found in the underlying Windom, but it is very rare
30 ABHighland Forest low dominance Ambocoelia samples (n = 7) Highland Forest Emanuella- Orbiculoidea dominated samples (n = 10)
Rhipidomella
Modiomorpha
Cypricardella
Greenops
Nuculites
P. constricta
Nuculoidea
Allanella
Tropidoleptus
Devonochonetes
Longispina
Mucrospirifer
Bembexia
Ontaria
Cyrtonella
Mediospirifer
Athyris
Emanuella
0 10203040 10 20 30 40 50 60 70 80 90 Percent Abundance Percent Occurrence Figure 6. Graphs of key Highland Forest taxa showing A) Each taxon's average percent abundance when it occurs in each group and B) Percentage of samples within each group that a taxon occurs in.
31 (Bembexia). What distinguishes the Emanuella-dominated samples are increased abundances and occurrences of a few key taxa in the Highland Forest group compared to the underlying
Windom, specifically Emanuella, Athyris, and Mucrospirifer. Interestingly, two Windom samples from HSTs (one sample each from the Gage Gully and Fisher Gully beds), one sample from a basal TST (and lowermost Amsdell bed), and one from an RST (Sheds bed) group more closely to Emanuella-dominated samples than to the ‘mainstream’ Windom gradient (Table 2), indicating that conditions were favorable at least a few times during ‘typical’ Windom conditions for these taxa to proliferate. Additionally, studies of Hamilton sequences undertaken stratigraphically below the current investigation have found Bembexia, Athyris, and
Mucrospirifer to be common components of Middle Devonian assemblages (Grasso, 1986;
Brower et al., 1978; Baird and Brett, 1983; Brett et al., 1990; Brower and Nye, 1991; Linsley et al., 1992; Newman et al., 1992) but they are not abundant in the short interval subsequent to the
Windom to Tully transition. Emanuella, on the other hand, is not found in the Hamilton Group, except for one unit within the lowermost Hamilton (Brower and Nye 1991). It is possible that this lower occurrence represents a different species of Emanuella (Cleland, 1903); regardless, it is unknown why this taxon is not found in any other intervening units.
Given that all the Highland Forest samples were deposited under relatively uniform conditions (TST to early HST), one might expect them to contain similar faunal assemblages, especially since many Windom taxa have been found to have wide ranges (Table 1). However, the two Highland Forest groups share only seven out of 19 genera (Fig. 6). Even among the seven shared taxa, abundances and occurrences are often different. For example,
Devonochonetes occurs in all samples of the low dominance cluster but only 20% of samples from Emanuella-dominated cluster in Figure 6B. Besides position within depositional sequence,
32
Sample Bed ST Emanuella Mucrospirifer abundance (%) abundance (%) TiloAms Lowest Amsdell TST 23 14 NLGage1A Gage Gully HST 0 56 BaFis1 Fisher Gully HST 80 0 ShetoShe Sheds RST 17 8
Table 2. Percentage of key taxa from the Emanuella-dominated assemblage that occur in non-boundary Hamilton samples.
33 what else could account for dissimilarities among these groups? While all of the Highland
Forest beds are silty shales, siltstones, or sandy siltstones, there are important lithologic differences between the two groups. Low dominance samples are shalier and more bioturbated as compared to Emanuella-dominated samples, which contain at least one of the minerals: pyrite, phosphate, and/or chamosite. Phosphate and chamosite typically form in times of extremely low to no sediment input. Thus, differences between the two faunas probably reflect differences in sedimentation and/or turbidity, and sample distribution along Axis 2 in Figure 4A may reflect these factors. As siliciclastic sedimentation began to wane, it is possible that the sea floor included sediment-starved patches that were colonized by a different assemblage (i.e.,
Emanuella-dominated samples). Three of the non-boundary Windom samples that group closely with the Emanuella-dominated samples (those in Table 2) are associated with widespread deepening and siliciclastic sediment reduction from sea level rise. Indeed, although many studies, including this one, observe the occurrence of Mucrospirifer in a wide variety of environments, several have highlighted its abundance in offshore areas with low sedimentation
(Brower et al., 1978; Brower, 1987; Savarese et al., 1986; Brower and Nye, 1991; Linsley,
1991). This accords well with the interpretation of reduced sedimentation in the environments representing the Emanuella-dominated deposits.
While DCA effectively distinguished samples from analogous parts of different systems tracts, it is not clear from Figure 4 if a consistent level of faunal fidelity is maintained within particular tracts of successive cycles, especially as the Windom/Tully boundary is approached.
To further investigate stratigraphic patterns, DCA sample scores from the stratigraphically most complete outcrop in the study, Highland Forest Park (Locality 10 in Fig.1), were graphed in stratigraphic order (Fig. 7). Stratigraphic spacing at this locality averaged less than 40 cm, with
34 Raw DCA scores 5-pt Moving Average DCA scores 2000 A B
1800
1600
1400
1200
4 Centimeters
1000 4
4 9
800 9
600 9
9 400 9
200
0 0 50 100 150 200 250 300 350 0 50 100 150 200 250 300 350 Axis 1
Deep Shallow Deep Shallow
TST Figure 7. DCA axis 1 scores plotted in stratigraphic order for RST locality 10. A) Unaveraged scores are coded according to position in parasequence. Samples with a '9' were taken from HST locality 9, '4' from locality 4. Star sample is a tie point to Low dominance Highland Forest connect localities. B) 5 point moving average scores. Emanuella dominated Highland Forest
35 the exception of two stratigraphic intervals: the 8 m regressive Taunton interval, which forms the
wall of a waterfall, and 2 m of the transgressive Spezzano bed, which was covered by stream
debris. These intervals were sampled at other nearby localities, and were “spliced” into Figure 7
using the distinct 40 cm South Lansing bed, sampled at all three localities, as a correlational tie
point. If anything, this substitution increases the potential to obscure meaningful patterns because
of possible differences among localities, and, thus, is a bias against observing a regular
stratigraphic pattern of repeated biofacies.
A raw data curve is presented in Figure 7A, while Figure 7B represents a five point
moving average. Small scale fluctuations are evident in Figure 7A and may represent local
vertical patchiness (sensu Webber 2002b), or perhaps slightly changing environmental
conditions. These minor variations are not unexpected, given the wide ranges of taxa and the
large amount of ordination space that each system tract occupies. Regardless of the cause, given
their limited magnitude, the focus of subsequent discussion will be on the smoothed curve (Fig.
7B) because it highlights the longer term stratigraphic signal.
Figure 8 shows the actual stratigraphy at the Highland Forest Park locality, plotted
against the five point moving average DCA scores. There is a strong correlation between DCA
Axis 1 sample scores and sequence tract interpretation. Moreover, this figure displays no distinction between tracts of successive cycles until the very end of Windom deposition. The two MFSs exhibit the smallest values, and the upper MFS (base of Gage Gully bed) exhibits a value that is almost identical to that of the lower MFS (base of the Fisher Gully bed). The HSTs show shallowing trends of similar magnitudes, and all RSTs display relatively constant values, with no obvious shallowing or deepening. As the Windom/Tully boundary is approached, the upper RST and TST of the Sheds bed deposits display a pattern that is similar, but not as large,
36 ully T
20 HST?
18 TST?
RST 16
Hamilton 14 HST Cycle 2 MFS
12 TST
10
8
6 RST Cycle 1
4 HST 2 MFS TST 0 RST Scale (m) 050 100 150 200 250 300 350 DCA Axis 1 sample scores
Deep Shallow
Figure 8. DCA axis one 5 point moving average scores plotted against the stratigraphic column at locality 10.
37 as the previous RST and TST. This may be a consequence of siliciclastic shutoff that occurred at this time (Heckel, 1973), or perhaps this was a lesser shallowing than previous RSTs and TSTs, and resulted in a slightly different assemblage.
Tully Samples
Figure 9 shows a DCA trial of the complete data set, including all Windom and Tully samples and taxa. In Figure 9A, the Windom gradient is still apparent, although more compressed and depicted along both Axes 1 and 2. In general, the Tully samples are more varied than those of the Windom, with some samples plotting along the Windom gradient and others that are distinctly separate from it. Figure 9B delineates taxa found only in the Windom, only in the Tully, or in both intervals. Several key Tully taxa are highlighted.
There are several distinct groups of Tully samples, each with unique faunal and lithologic characteristics. The lowermost Tully samples are interpreted as relatively deep water deposits, and are particularly interesting because they form three discrete groups (Fig. 9A). The most striking of these is the group of samples (labeled 'Rhyssochonetes-dominated') that score most highly on Axis 1, and is discrete from all other samples. These samples are overwhelmingly dominated by the brachiopod Rhyssochoenetes aurora (Figs. 9B and 10), and many samples contain only Rhyssochoenetes. Like the Eumetabolotoechia-dominated Windom samples, the small number of individuals (11) in these samples faithfully represents the monospecific nature of these deposits, as additional specimens were counted in the field and no new taxa were encountered. The bulk of these samples are from the two lowermost Tully units, the DeRuyter and Cuyler beds (Fig. 11), in which extensive R. aurora pavements are common. A few samples are from the less intensively sampled upper Lower Tully. Lithologically, these calcareous
38 DCA of All Samples 500 A 450
400
350
300
250
200 upper Tully West Brook Shale samples 150 lower Tully Eumetabolotoechia- dominated samples 100 lower Tully Emanuella- dominated samples lower Tully Rhyssochoetes- 50 dominated samples Windom samples 0
Axis 2 DCA of All Taxa 500 B 450 Emanuella 400 Mucrospirifer Hypothyridina Strophomenid "A" 350
300 Atrypid fragment 250 Rhyssochonetes Strophomenid fragment 200 Indet. Rhynchonellid
150
100 Eumetabolotoecia Windom only occurrence Tully only occurrence 50 Windom & Tully
0 0 100 200 Axis 1 300 400 500 600 Figure 9. DCA axis 1 and 2 scores plotted for all samples (A) and taxa (B). Key taxa are highlighted in B.
39 A Tully Eumetabolotoechia- B dominated (n = 4) Rugosa Tully Emanuella- dominated (n = 14) Allanella Tully Rhyssochonetes- dominated (n = 15) Tropidoleptus
Emanuella
Mucrospirifer
P. emarginata
Eumetabolotoechia
Longispina
Indet. Crinoid
Ambocoelia
Devonochonetes
Hypothyridina
Strophomenid 'A'
Rhynchonellid
Pseudoatrypa
Eoschuchertella
Rhyssochonetes
01020304050607080 0102030405060708090100 Percent Abundance Percent Occurrence Figure 10. Graphs of key Tully taxa showing A) Each taxon's average abundance when it occurs in each Lower Tully group and B) Percentage of samples within each group that a taxon occurs in.
40 41 Starred taxaoccurinboth theW the num Figure 12.Percentoccurrenceofthem
ber ofsa
m *Devonochonetes *
p *Paleoneilo constricta* les inwhichataxonoccurs, *Mucrospirifer* *Tropidoleptus* *Eumetabolatoechia* *Nuculoidea* i ndom andTully *Longispina* *Indet Crinoid*
o *Allanella* st commonta *Emanuella* *Clam fragment*
divided bythetotalnumber ofsam *Paleoneilo emarginata* Fenestrate
xa withinthestudied Nuculites Spyroceras Mediospirifer *Indet/Frag Trilobite* *Greenops* *Rhipidomella* *Spirifer fragment*
interval. Occurrence= Pustulatia pustulosa Rhyssochonetes aurora Stictopora p l
e Lingula s (152). 0% 5% 10% 15% 20% 25% 30% 35% 40% 45% 50% Percent Occurrence 42 siltstones are carbonate-enriched compared to the underlying Windom; the DeRuyter has an average carbonate content of 40% (Heckel, 1973). The Cuyler Bed is not as calcareous, and has an average carbonate content of 23% (Heckel, 1973). R. aurora was not found in the sampled units of the Windom during this project, nor has it been reported from other New York Hamilton studies, and can thus be considered a taxon unique to the Tully Formation. The brachiopod
Hypothyridina and, possibly, a new rhynchonellid species are found in these samples, and are also unknown in Hamilton deposits. Other 'accessory' taxa found at low abundance in this group are among the most common and eurytopic Middle Devonian fossils of New York, namely:
Tropidoleptus, Mucrospirifer, Paleoneilo, Longispina, Ambocoelia, and Devonochonetes (see
Figs. 5, 8, 12 and Savarese et al., 1986; Brett et al., 1990; Brower and Nye, 1991; Linsley, 1991;
Linsley et al., 1992; Newman et al., 1992; Laffery et al., 1994). Additional uncommon elements of these samples, Pseudoatrypa and Eoschuchertella, were both found in Windom deposits during this study, but at low abundances (about 6% for each taxon), whereas other studies of
Hamilton biotas have found these taxa as common or abundant constituents of assemblages
(Koch, 1981; Baird and Brett, 1983; Grasso, 1986; Brett et al., 1983; Brett et al., 1990). Thus, the dominance of one genus, Rhyssochonetes, is fundamental in differentiating this assemblage.
Another cluster of Tully samples score most highly on Axis 2, and are labeled
'Emanuella-dominated' in Figure 9A. Common Windom taxa are present in low abundances, similar to Windom taxa abundances in samples of the Rhyssochonetes-dominated group.
Emanuella-dominated Tully samples intermingle with those of the Highland Forest Emanuella- dominated group, and thus, contain an assemblage that passed through the Windom to Tully transition with only moderate change. About 40% of the total, combined pool of taxa is shared between these two groups, and about 66% of the Emanuella-dominated Tully fauna is composed
43 of taxa also recognized in the Emanuella-dominated Highland Forest samples. The high abundance and occurrence of Emanuella, and to a lesser extent Mucrospirifer, are the main taxonomic factors that group these samples together (compare the Emanuella-dominated groups of Figs. 6 and 10).
The Emanuella-dominated Tully samples are mainly from the DeRuyter and Cuyler beds, but are lithologically distinct from the Rhyssochonetes-dominated samples, which were collected from the same beds. Emanuella-dominated samples come from siltstones and sandstones that contain pyrite, phosphate, and/or chamosite. This lithology is virtually identical to that of the
Emanuella-dominated Highland Forest samples, and thus, key taxa probably persisted in the environment that this lithology represents through time. Both the Emanuella-dominated and
Rhyssochonetes-dominated assemblages are found in the upper Lower Tully, indicating that the two faunas, although coeval, remained distinct through time (Fig. 11).
Four additional lower Tully samples group in the lower lefthand corner of the DCA graph
(Fig. 9A) as they are dominated overwhelmingly by the brachiopod Eumetabolotoechia (Figs. 9B and 10). The few other taxa that occur in these samples are also found in Windom and other
Lower Tully deposits. Lithologically, these samples are somewhat similar to those of the
Emanuella-dominated samples in that they are chamositic siltstones, but contain a larger component of sand-sized grains and mica (personal observation; Smyrna Bed of Heckel, 1973), which was not found in Emanuella-dominated samples. These samples were collected from a locality that likely records the deepest lower Tully environments, as it is situated within the sub- basin (easternmost samples on Fig. 11) (Heckel, 1973). Eumetabolotoechia is most dominant in dysoxic environments, both in this study (Fig. 5) and throughout the Devonian (Kammer et al.,
1986; Thompson and Newton, 1987; Vogel et al., 1987), and thus, these Tully samples may have
44 been deposited under dysoxic conditions. This assemblage was found in one sample of the lowermost Tully (DeRuyter Bed) and three samples within the upper Lower Tully, indicating that it persisted through that time (Fig. 11).
Figure 11 shows a geographic separation of Lower Tully sample groups, with
Rhyssochonetes-dominated samples occurring to the west, Emanuella-dominated samples more common to the east, and Eumetabolotoechia dominated samples at easternmost localities. This separation of faunas may be related to paleo-depth, with Eumetabolotoechia samples deposited in the deepest environments, Emanuella samples slightly shallower, and Rhyssochonetes samples slightly shallower still. Alternatively, faunas may be responding to siliciclastic input, with siliciclastic dominated environments in the east and more sediment starved, mixed to carbonate dominated environments in the west. Rhyssochonetes, Eumetabolotoechia, Hypothyridina, and
Emanuella have all been reported in the Middle and Upper Tully, stratigrapically above the units studied in this project (Heckel, 1973; Baird and Brett, 2003), although it is unknown whether the faunal associations discussed here persisted as well.
The final group of Tully samples was collected not from the Lower Tully, but from the
West Brook Shale Bed, a unit within the Upper Tully, and the stratigraphically highest bed studied (Fig. 2). In Figure 9, these four samples group with the highest scoring Windom samples on Axis 1, and contain a similar high diversity, low dominance assemblage composed of corals, bryozoans, and many brachiopod genera. While all of the studied lower Tully units contained a variety of calcareous siltstones or sandstones, the younger West Brook Shale bed is a calcareous mudstone, and is lithologically similar to shallow water Windom deposits. This implies that
Windom faunas migrated out of the studied area at the onset of Tully deposition, when siliciclastic production waned, and that many of the same taxa later returned when more
45 favorable habitats again appeared during the deposition of the West Brook Shale (see Bonelli et al., 2003, for additional discussion). Because this bed is significantly above the Windom/Tully boundary, it was not included in further analyses.
Summary of Results
Within a geologically short interval of time (probably much less than 0.9 ma, House,
1995) a faunal gradient recurs, with moderate variation, within successive small-scale cycles of the upper Windom Shale in New York State (Figs. 8 and 11). Detrended Correspondence
Analysis (DCA) arrayed fossil samples of the uppermost Windom along axes that are inferred to represent physical variables correlated to paleo-depth. Position along Axis 1 correlates well to position within fourth order cycles, with deepest water assemblages found in early highstand deposits and shallowest water associations found at the bases of transgressions. Stratigraphically higher samples, near the Windom/Tully boundary (Sheds through Highland Forest interval) are inferred regressive through highstand deposits that, while comparable in composition, consistently fail to score as highly on DCA Axis 1 as samples of previous RST, TST, and HST deposits (Fig. 8). Therefore, there was a slight, but noticeable, change in biotic composition as the boundary is approached. Moreover, at this time, a fauna ('Emanuella-dominated Highland
Forest' Fig. 4) composed of taxa that were otherwise rare within the studied interval became prevalent in a previously uncommon lithofacies (pyritic/phosphatic siltstone). This fauna was coeval with a biota more typical of Windom assemblages ('low Dominance Highland Forest' Fig.
4), but these two associations remained compositionally distinct (Fig. 6). The more typical
Windom fauna did not persist across the boundary, but most of the major elements of the
Emanuella-dominated assemblage remained in their preferred environment into the Lower Tully.
46 This fauna is then found with two coeval, but distinct assemblages; the Rhyssochonetes-
dominated assemblage in calcareous siltstones, and the Eumetabolotoechia-dominated
assemblage in chamositic/micaeous sandstones (Fig. 11). The most abundant taxa of these
assemblages, Emanuella and Eumetabolotoechia, were also found in Hamilton deposits in this
study, whereas Rhyssochonetes and Hypothyridina appear to have their first occurrences in the
Appalachian Basin at the lowermost Tully boundary. All three of these coeval assemblages
remain distinct throughout the Lower and lower Middle Tully (Fig. 11). Overall, therefore,
faunal patterns were complex immediately prior to and following the Hamilton/Tully boundary.
Implications
Analyses of faunal dynamics at this regional turnover boundary have revealed that
turnover was likely caused by physical changes near the Windom/Tully boundary, as lithological
changes record probable environmental transitions and correlate strongly with biotic changes.
The uppermost Windom low dominance or ‘typical’ Windom assemblage does not persist into the lowermost Tully, while the uppermost Windom Emanuella-dominated assemblage does. It
appears that these faunal changes were associated with siliciclastic shutoff and related
development of phosphate and chamosite, and/or the onset of carbonate production, and thus,
taxa appear to be responding to changes in the physical conditions of the basin (sensu Bambach,
1994; Miller, 1997). In general, turnover at the Windom/Tully boundary is characterized more
by changes in relative taxonomic abundance, rather than by the introduction of 'exotic' biotas,
implying that taxon abundance information is required to fully understand regional biotic
turnovers. This corroborates previous ecologic studies that have cited the importance of
collecting abundance data (e.g., see Jackson, 1994).
47 It is interesting that both the Emanuella-dominated and Eumetabolotoechia-dominated assemblages are found in both the Windom and Tully, and that only a handful of Tully taxa are not found within the Hamilton (Figs. 9 and 12) suggesting that the biotic transition in this case was not atypical of what one would simply expect as a consequence of turnover at a sequence boundary (e.g. the transition from Windom Cycle 1 to Cycle 2 in Figure 11). Given that this was an unusual perturbation at the Windom/Tully boundary, it might be tempting to therefore argue that taxa withstood it because assemblages were ' buffered’ by some form of biotic interactions
(Morris et al. 1996; see discussions within Miller, 1996 and Ivany, 1996, 1999). However, there is compelling evidence to suggest that instead, this pattern reflects the component taxa's individual ecologies. Taxa that persisted into the Tully are documented to have broad environmental tolerances, both within the studied interval and throughout the Devonian
Appalachian Basin (Figs 5, 8, and 12), and thus it is not surprising that they persisted across the boundary.
A possible exception to this is the genus Emanuella, which is found in the lowermost
formation of the Hamilton (Marcellus Formation) and then is absent until the studied interval, an
almost 4 million year hiatus (Brett and Baird, 1994). Although it is unknown why Emanuella is
absent from the intervening units, it is clear it must have been somewhere else. Within the
studied interval, Emanuella is rare preceding the boundary interval and then becomes abundant
throughout the transition. Within the Marcellus, Emanuella occurs within ‘dark carbonaceous
shale’ (Brower and Nye, 1991). Peak abundance of Emanuella occurs within the uppermost
Hamilton (Highland Forest bed) and lowermost Tully (DeRuyter Bed), deposits that are
interpreted as sea level highs with reduced siliciclastic input, and so, perhaps, Emanuella could
not tolerate earlier conditions of siliciclastic sedimentation within the intervening units of the
48 Hamilton Group. Additionally, the abundance of Emanuella varies widely within the few samples it occurs in preceding the boundary interval (Fig. 5 and Table 2), suggesting that it was not part of a biologically integrated community (e.g. Elton, 1927). Although only one sample of the Amsdell Bed from this study was highly dominated by Emanuella, other studies have found
Emanuella to be profuse in this unit in western New York, near Buffalo (Brett and Baird, 1996), so perhaps its abundance varied dramatically across the basin as well as through time.
It would be beneficial to also make detailed comparisons between the Hamilton
Eumetabolotoechia dominated samples and those of the Tully. Unfortunately, this biofacies was only found within four samples of the Tully, making detailed comparisons difficult. It is clear that Eumetabolotoechia was overwhelmingly abundant in dysoxic environments both before and after the turnover, and appears to have been a eurytopic taxon.
Figure 4 illustrates Windom samples arrayed along a continuous gradient with no compositional breaks between recognized groups. Clearly, many of the studied taxa exhibited overlapping paleoenvironmental ranges, supporting the idea of little biotic integration among taxa within assemblages. Even within the Tully, where three different ‘biofacies’ are recognized, the majority of taxa are shared, and the abundance of only one or two key taxa distinguishes assemblages from one another (Fig. 10). It is difficult to envision how each of these
‘communities’ could have a high degree of biotic interaction if many taxa are shared across assemblages (Fig. 12).
Another interesting, related aspect is the relationship between the Emanuella-dominated assemblage and the Windom gradient (Fig. 4). Emanuella-dominated samples contain many of the same taxa as the typical Windom highstand deposits, and intermingle somewhat on a compositional basis with the Windom gradient, rather than grouping discretely from it.
49 Emanuella-dominated samples appear to come from an environment that was more sediment starved, but otherwise similar to, ‘typical’ Windom deep water, dysoxic environments, as
Emanuella-dominated samples from both the Windom and Tully are associated with periods of sea-level highstand and siliciclastic sediment reduction. This intermingling of the Emanuella- dominated assemblage and the Windom gradient also argues for low biotic interaction.
This returns to one of the main questions of this study, was turnover abrupt? The most appropriate answer is that in some ways yes, it was, and in other ways, no, it was not. The
Eumetabolotoechia assemblage apparently remained unchanged after the turnover event. The
Emanuella assemblage, rare within the studied interval, became abundant and widespread during the highest Windom and remained so within the Tully, suggesting a gradual pattern of buildup.
The Rhyssochonetes assemblage is unique to the Tully and could represent an example of an abrupt incursion. Some caveats are necessary, however. It is possible, due to both nondeposition at the Windom/Tully boundary and the restricted geographic conformable area, that the abrupt appearance of Rhyssochonetes is artifactually enhanced (Holland, 1995, 1996).
While it is probable that this disconformity is minor (Baird and Brett, 2003) it will be difficult to extend the range of study geographically, as the contact between the Windom and Tully becomes more disconformable westward. It may be possible to sample eastward, although sedimentation increases dramatically because of the Catskill Delta, and thus correlation becomes difficult.
Therefore, while the appearance of Rhyssochonetes may have been abrupt, it is impossible to say definitively that this was the case.
It is clear that turnover was not synchronous across the boundary, as the Emanuella assemblage is seen both before and after the transition interval, while the Rhyssochonetes assemblage is found only within the Tully. Even if the abrupt appearance of Rhyssochonetes is
50 artifactually sharpened, Emanuella still occurs below the boundary samples of the Windom (Fig.
11), and therefore still appears earlier.
Why does the lowermost Tully unit (DeRuyter) appear to be more heterogeneous faunally
(i.e., exhibit assemblages that are much more compositionally variable), than any of the underlying Windom beds, which are known to exhibit faunal gradients (Lafferty et al., 1994;
Bezusko, 2001; Bonelli et al, 2003)? In part this may be a sampling issue – the DeRuyter was extensively sampled laterally, much more so than any Windom bed. However, it is unlikely that sampling provides the sole explanation. Low sedimentation, coupled with the switch from a siliciclastic to mixed system may have caused the development of patches on the Tully sea floor.
While all Tully samples have a silt (and in some cases sand) component, mineral composition varies from chamosite, to phosphate and pyrite, to carbonate enriched siltstones. Indeed, during the beginning of siliciclastic shut-off, within the Highland Forest bed, two distinct coeval biotas are observed. Thus, it seems reasonable that the seafloor of the Appalachian Basin became more heterogeneous as turnover progressed, and that different patches were occupied by different assemblages.
The faunal complexity found at this boundary was unexpected at the outset of this project, and could have gone unnoticed if not for detailed sampling of stratigraphically complete outcrops. Perhaps turnovers in general are more complex than previously assumed.
51 CONCLUSIONS
1) A depth controlled gradient existed within the uppermost Windom Shale of the Appalachian
Basin. This interpretation is supported by independent lithologic and sequence stratigraphic evidence, and agrees well with previously published depth estimates for certain taxa. In
Detrended Correspondence Analyses (DCA) of fossil samples, the Windom gradient is expressed primarily along DCA Axis 1, which has been found to be a common proxy for depth correlated variables in previous ecologic and paleoecologic studies.
2) The Windom gradient is composed of wide ranging taxa whose abundances, rather than occurrences, vary systematically along the gradient. A lack of biotic interaction is demonstrated by the abundance of taxa with broad environmental ranges that occur in many samples and environments.
3) Successive parts of systems tracts display only moderate compositional variability until just prior to the Windom/Tully boundary. Within the uppermost Windom, a formerly uncommon fauna (Emanuella-dominated assemblage) is found within a previously unrecorded facies. This fauna was coeval, but distinct, from assemblages more typical of the Windom. Within the Tully, two assemblages documented from the Windom (Emanuella-dominated and Eumetabolotoechia- dominated assemblages) and one assemblage unique to the Tully (Rhyssochonetes-dominated assemblage) were found. These observations imply that turnover was complex, as the
Rhyssochonetes-dominated assemblage appeared abruptly within the Tully, the Emanuella-
52 dominated assemblage exhibited a pattern of more gradual buildup, and the Eumetabolotoechia- dominated assemblage appears to have been unaffected by turnover.
4) Biotic turnover was asynchronous, in that two assemblages occur in both the Windom and
Tully, while a distinct Tully fauna does not appear until Tully deposition begins.
5) With the exception of one or two important taxa (i.e. Rhyssochonetes and Hypothyridina)
Tully biotas are distinct from those of the Windom because of changes in the abundances of key taxa, rather than their wholesale replacement. This highlights the importance of collecting abundance data for paleoecologic analyses within the studied area.
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Van Wagoner, J. C., Posamentier, H. W., Mitchum, R. M. J., Vail, P. R., Sarg, J. F., Loutit, T. S., and Hardenbol, J., 1988, An overview of the fundamentals of sequence stratigraphy and key definitions: in Wilgus, C. K., Hastings, B. S., Kendall, C. G. S. C., Posamentier, H. W., Ross, C. A., and Van Wagoner, J. (eds.), Sea-level changes: An integrated approach: v. 42: Society of Economic Paleontologists and Mineralogists Special Publication, p. 39- 45.
Van Wagoner, J. C., Campion, K. M., Mitchum, R. M., and Rahmanian, V. D., 1990, Siliciclastic sequence stratigraphy in well logs, cores, and outcrops; concepts for high-resolution correlation of time and facies, Methods in Exploration Series: v. 7: American Association of Petroleum Geologists, p. 55.
61 Vogel, K., Brett, C. E., and Golubic, S., 1987, Endolith associations and their relation to facies distribution in the Middle Devonian of New York State, U.S.A: Lethaia, v. 20, p. 263- 290.
Webber, A. J., 2002a, High-resolution faunal gradient analysis and an assessment of the causes of meter-scale cyclicity in the type Cincinnatian Series (Upper Ordovician): PALAIOS, v. 17, p. 545-555.
Webber, A. J., 2002b, The effects of faunal patchiness on the use of gradient analysis for regional, high-resolution correlation in the type Cincinnatian Series (Upper Ordovician): Geological Society of America, Abstracts with Programs, v. 34, p. 35.
Whittaker, R. H., 1975, Communities and ecosystems, Second edition: Macmillan, New York, 385 p.
62 APPENDIX I.
Locality register and sample position.
63 Loc # Samp Bed Locality information Name 1 KasSpez1 lowest Spezzano, creek level Kashong Glen, town of Bellona KasSpez2 Spezzano 1m above KasSpez1 KasSpez3 Spezzano 1m above KasSpez2 KasKaBd Karla bed 1m above KasSpez3 KasGage Gage Gully 1m above KasKaBd MineWBS West Brook Shale Mineagar Brook, 2DeRySkanbasal DeRuyter Bloomer Creek, town of Skaneateles BoGag1 Gage Gully directly beneath the Tully BoGag2 Gage Gully 0.6m beneath Tully BoSpez1 Spezzano 1.34m beneath Tully BoSpez2 Spezzano 2.24m beneath Tully BoTau1 Taunton 0.9m below BoSpez2 BoTau2 Taunton 1m below BoTau1 BoTau3 Taunton 1.3m below BoTau2 BoTau4 Taunton 1.m below BoTau3 BoTau5 Taunton 1.m below BoTau4 3 BaFis1 Fishers Gully ground level Barnum Shale Pit, town of Skaneateles BaFis2 Fishers Gully 1.5m above BaFis1 BaFis3A Fishers Gully 1m above BaFis2 BaFis3B Fishers 0.92m below BaFis/Ta BaFis4 Fishers 0.55m above BaFis3B BaFis/Ta Taunton/Fishers cont 0.37m above BaFis4 4 GroLans Lansing top Groves creek, town of Skaneateles GroSpez1 Spezzano 0.41m above GroLans GroSpez2 Spezzano 0.4m above GroSpez1 GroGage Gage Gully 0.15m below Tully 5 FaBrFB Fall Brook Bed, 1cm thick Fall Brook, town of Skaneateles FaBrGag Gage Gully FaBrLans Lansing top 6 CaTau1 Taunton 1m below first silt ledge Carrs Quarry, town of Tully CaTau2 Taunton 0.5m below first silt ledge CaTau3 Taunton first silt ledge CaTau4 Taunton 0.5m above first silt ledge CaTau5 Taunton 1m above first silt ledge CaSh1 Sheds 1.5m below DeRuyter CaSh2 Sheds .75m below DeRuyter CaDeRuy DeRuyter CaCuylr Culyer Bed 0.56m above base DeRuyter CaFab? Fabius?/Cuyler 0.95m above CaCuylr CaTulVa Tully Valley 2m above CaFab? CaShenoHF top Sheds directly below DeRuyter 7 JuHiF/Sh top Sheds/Highland Forest contact Junes Ravine, town of Tully, Heckel loc. 1CD DeRyJune DeRuyter HiFoJune uppermost Highland Forest ShedJune Sheds 8 in below Tully 8 ShedSFab Sheds 5-10 in below Tully creek S of Fabius 9 TiSHnoHF Sheds 0.95m below DeRuyter Tinkers Falls county park, town of Fabius TiSheds Sheds 1m below TiSHnoHF TiGage1 Gage 2.2m below TiSheds TiGa/Sh Gage/Sheds transition 1m above TiGage1
64 Locality Samp Bed Locality information # Name 9 TibaGag Gage 0.1m above TiSpezz2 TiSpezz2 Spezzano 0.1m below TibaGag TiSpezz Spezzano 0.45cm above TiSpezz2 TiLans Lansing 1.1m below TiSpez TiTau1 Taunton 1.7m below TiLans TiTau2 Taunton 2m below TiLans TiTau3 Taunton 3.3m below TiLans TiTau4 Taunton 5m belowTiLans TiTau5 Taunton(?) 6.4m below TiLans TiFis2A Fishers Gully 2.2m above TitoAms TiFis2B Fisher Gully ~2m laterally from TiFis2A TiFi1 Fishers Gully 0.87m above TitoAms TitoAms top Amsdell TiloAms Base Amsdell 10 HFBS1 Bear Swamp 0.5m below HFAmsLS Highland County Park, town of Fabius HFBS2 Bear Swamp 0.37m below HFAmsLS1 HFAmsLS1 lowest Amsdell Bed HFAmsSH2 Amsdell 0.08m above HSAmsLS1 HFAmsdH3 Amsdell 0.19m above HSAmsSH2 HFAmsLS4 Amsdell 0.08m above HSAmsSh3 HFsitaAm siltstone 0.07m above HFAmsLS4 HFFG1 Fishers Gully 0.19m above HFSitaAm HFFG2 Fishers Gully 0.19m above HFFG1 HFFG3 Fishers Gully 0.17m above HFFG2 HFFG4 Fishers Gully 0.05m above HFFG3 HFFG5 Fishers Gully 0.30m above HFFG4 HFFG6 Fishers Gully 0.24m above HFFG5 HFFG7 Fishers Gully 0.24m above HFFG6 HFFG8 Fishers Gully 0.27m above HFFG7 HFFaBrk Fall Brook Bed 0.1m above HFFG8 HFTauton Taunton 0.18m above HFFaBrk HFLansng Lansing HFKarbd1 Karla 0.15m above HFLansng HFKarbd2 Karla 0.10m above HFKarbd1 HFGage1A Gage Gully 0.30m above HFKarbd2 HFGage1B Gage Gully 0.50m laterally from HFGage1A HFGage2 Gage Gully 0.36m from HFGage1B HFGage3 Gage Gully 0.14m from HFGage2 HFGage4 Gage Gully 0.24m from HFGage3 HFGage5 Gage Gully 0.38m from HFGage4 HFbasShd Sheds 0.12m above HFGage5 HFShed1 Sheds 0.33m above HFbasShd HFShed2 Sheds 0.42m above HFShed1 HFShed3 Sheds 0.40m above HFShed2 HFShed4 Sheds 0.44m above HFShed3 HFShed5 Sheds 0.42m above HFShed4 HFShed6 Sheds 0.50m above HFShed5 HFShed7A Sheds 0.27m above HFShed6 HFShed7B Sheds 1m laterally from HFShed7A HiFor1 Highland Forest 0.31m above HFShed7B
65 Locality Samp Bed Locality information # Name HiFor2 Highland Forest 0.40m above HiFor1 Hifor3 Highland Forest 0.50m above HiFor2 10 HiFor4 Highland Forest 0.43m above HiFor3 HiFor5 Highland Forest 0.18m above HiFor4 HiFor6 Highland Forest 0.13m above HiFor5 HiFor7 Highland Forest 0.38m above HiFor6 HiFor8 Highland Forest 0.24m above HiFor7 HiFor9 Highland Forest 0.11m above HiFor8 HFbasDeR DeRuyter 0.50m above HiFor9 HFupDeRy DeRuyter 0.16m above HFbasDeR HFCuyl1 Culyer 0.45m above HFupDeR HFCuyl2 Cuyler 1.14m above HFCuy1 HFCuyl3 Cuyler 0.26m above HFCuy2 HFFabi Fabius?/Cuyler 0.48m above HFCuy3 11 CaRDHiFo Highland Forest beneath Tully Carey Rd, DeRuyter 12 ShetoShe top Sheds creek 3.5 mi NE of Sheds NEShHiFo Highland Forest directly beneath Tully 13 XRdtHiFo uppermost Highland Forest Carpenters Rd SW of Sheds 14 DeSSheds DeRuyter creek S of Sheds subDRSSd Highland Forest directly beneath Tully WBSSofSherWest Brook Shale 15 DeRtoShd top Sheds Quarry northwest of DeRuyter, Heckel loc. 3A DeRbHiFo base Highland Forest baDeNWDR base DeRuyter 16 WBS PVT West Brook Shale road/creek 0.6 mi south of the town DeRuyter baTuSDR1 basal DeRuyter DeRySDer DeRuyter baTuSDR2 basal DeRuyter 17 WBSWerns West Brook Shale Werners Glen, Georgetown ShedWern Sheds 18 HoGlNeLi uppermost Highland Forest Houghtlings Glen, town of Edmeston baTuHoGl basal Tully 19 Gag1walt lowest Gage Gully Walt Phillips Rd. town of New Berlin Gag2walt ~1 meter above Gage1walt sample Gag3walt ~1 meter above Gage1walt sample GageGren Gage Gully Greens Gulf, New Berlin 20 baTuSPit basal Tully creek 0.7 mi S of Pittsfield 21 upNeLsNL upper New Lisbon Town of New Lisbon baNeLsNL ~5 feet above basal New Lisbon baLaNwLs basal Laurens/top New Lisbon/ Tully NLGage1 Gage Gully NLGage1A Gage Gully 1m laterally from NLGage 1 NLLansing Lansing NLDRuyt1 basal DeRuyter NLDRuyt2 DeRuyter shell bed 1 NLShelB2 top of New Lisbon, shell bed 2 NLSB3Fab Shell bed 3 Fabius NLSB4Lau Shell bed 4 Laurens NLCorazo ~7 feet above NLSB5Lau NLSB5Lau Shell bed 5 upper Laurens NwLsNENL New Lisbon member/base Tully creek 1 mi northeast of New Lisbon baTuNENL basal Tully
66 APPENDIX II.
Middle Devonian uppermost Hamilton Group and Tully Formation field census raw data. Sample and locality codes correspond to those in Appendix I. Total matrix is 152 samples and 111 taxonomic categories.
67
HFsitaA HFAm HFAm HFAm HFAm H H C C C C C C C C C C C C T T T T T T T T T T T T T T T T T T N N N N N N N N N N i i i i i i i i i i i i i i i i i i a a a a a a a a a a a a F F L L L L L L L L L L l t F F F T T T T T L S K b G G S ShnoH S T F C D S S T T T T T o o B B S C S S S D D L G G a i i i p h a a a a a a a a a h a h h A A 1 s s u a a a a a u e G B B B h a u u u u u n S S e e o R R a a r / g 2 2 b u u u u u e 2 1 l R y S b n e m m z d 5 4 3 2 1 s g g 2 1 r V 5 4 3 a e n ? B A 5 4 3 2 1 u u sdH3 sL sS sL l a l z d h s s u e e g 1 L L F r B o s s y y a z i m 1 1 y n a H a a F H2 t t o S4 S1 2 A 2 1 b g u u F 0 007 006 000 0 000 00 0 001 0 000 001 0 006 020 000 000 000 002 000 000 0 0 03 400 285 332 200 000 000 0 720 102 002 004 024 03 100 000 000 000 000 000 00 009 101 0 Ambocoelia 2 1 1 0 46 36 1 40 3 1 0 1 Emanuella 1 1 1 1 15 0 3 0 0 0 3 1 0 1 Mucrospirifer 0000 1000 0300 0101 0000 0000 0100 0000 0000 0000 0 0000 0000 0000 0700 0000 8205 0300 1000 0000 0200 2000 0000 0010 0000 0000 0000 0004 0000 0000 9000 0000 0000 0000 0601 0200 0100 0007 0101 0000 0000 0000 0000 0010 0000 4400 9000 Mediospirifer 1
000 Allanella Cyrtina Pustulatia 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 101 000 000 000 000 000 000 010 000 600 000 010 000 000 000 000 000 000 000 000 000 111 000 000 000 000 000 000 000 000 000 Pseudoatrypa Spinatrypa Elita fimbriata 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 1000 0000 0000 0000 0000 0010 0000 3000 0000 0010 0000 0000 0000 0000 0000 2000 0000 0000 0000 0000 0000 0000 0000 0000 0600 0000 0000 0000 0000 0000 0000 Megastrophia Mesoleptostrophia Protoleptostrophia Trematospira 0800 0800 1000 0640 0000 0000 0320 00 0000 0000 02 0340 0000 0000 013 0200 0020 0000 0 0 0200 0110 0000 0000 0000 000 0000 040 0220 0300 0310 0020 0010 0000 0109 0120 0100 0000 0 0 0110 0100 0820 0500 0 0410 0010 Strophodonta 1 1 1 1 1 000 400 030 100 230 Devonochonetes 1 2 2 6 Longispina 2 1 1 0 0 4 9 2 Rhyssochonetes 0000 0000 0000 0000 0000 0000 0000 0000 000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 3000 0000 1000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 Eoschuchertella Douvillina Retichonetes 1
0 Productids 0100 0000 0000 0300 0080 0 0 0000 0000 0000 00 0200 02 0080 0700 0019 00 0300 0030 1500 00 0080 04 0000 3100 00 0300 0000 1100 3200 00 16 0000 0020 0200 00 0302 0100 1000 0300 0200 0500 0020 0000 0000 0000 0000 Rhipidomella 1 1
300 200 Tropidoleptus 1 2 1 1 1 1 1 2 2 7 4 1 3 0 1 1 0 2 Eumetabolotoechia 0 0 1 0 0 0 0 0 0 Hypothyridina 00 00 00 00 00 00 00 00 0 00 0 00 0 00 0 00 0 00 00 0 00 00 00 00 00 20 01 0 0 00 0 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 Cupularostrum 0 0 0 0 0 0 0 0 0 Rhynchonellid
68
HFsitaA HFAm HFAm HFAm HFAm H H C C C C C C C C C C C C T T T T T T T T T T T T T T T T T T N N N N N N N N N N i i i i i i i i i i i i i i i i i i a a a a a a a a a a a a F F L L L L L L L L L L l t F F F T T T T T L S K b G G S ShnoH S T F C D S S T T T T T o o B B S C S S S D D L G G a i i i p h a a a a a a a a a h a h h A A 1 s s u a a a a a u e G B B B h a u u u u u n S S e e o R R a a r / g 2 2 b u u u u u e 2 1 l R y S b n e m m z d 5 4 3 2 1 s g g 2 1 r V 5 4 3 a e n ? B A 5 4 3 2 1 u u sdH3 sL sS sL l a l z d h s s u e e g 1 L L F r B o s s y y a z i m 1 1 y n a H a a F H2 t t o S4 S1 2 A 2 1 b g u u F 0 000 000 000 000 000 000 0 000 000 000 000 000 000 000 000 000 000 000 000 000 2 1 0 000 000 000 200 000 000 000 0 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 Athyris 0 0 0 0 0 0 Megakozlowskiella 0 0 0 0 0 0 "Orthis" lepidus 0000 0000 0000 0000 10 0000 0000 0000 0000 0000 0000 0000 0000 0081 0000 0000 1000 0000 0000 0000 0000 0000 0100 0000 0000 0000 0000 0000 0000 0000 1000 1000 0000 0000 0010 0100 0000 0000 0001 0000 0000 1020 0000 0000 0000 0000 0000 Lingula Crania 1
5 Orbiculoidea
1 Lidstroemella 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 001 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 Pholidops Unkn Inarticulata Cranaena 1020 0003 0010 0000 0001 0000 0000 0000 0011 0010 0010 0000 0000 0000 0000 0003 0101 0000 0000 0000 0000 2010 0000 0000 0000 0000 0001 0000 0000 0000 0000 000 0001 0000 0000 0000 0000 0010 0000 0000 0000 0000 0002 0011 0000 0022 0000 Cryptonella Unkn Terebratulid Nuculoidea 1
1 Nuculites 0102 0000 0002 0000 0012 0000 0000 0000 0000 0001 0001 0000 0000 0000 0000 0006 0005 0000 0000 0000 0000 0102 0000 0000 0000 0006 0009 0000 0000 0001 0002 0002 0003 0000 0006 1014 0003 0002 0001 0001 000 0005 0008 0105 0002 0004 0003 Nuclea Modiomorpha Modiella 1
5 P. constricta 0000 0000 0000 0000 4200 0000 0000 0000 0000 0000 1000 0000 0000 0000 0000 1000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 5000 0000 0000 0100 0000 1000 4000 1000 0000 1000 0000 0000 0000 0020 0000 1010 1000 0000 3000 0000 0000 P. emarginata P. filosa Pholidella Cimitaria 0000 1000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 1000 0000 0000 0000 0000 0000 0100 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0011 0000 0000 0100 0000 0000 0000 0000 Tellinopsis Cypricardella Phthonia Meristella 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 01 00 00 00 00 00 00 00 Mytlops Ontaria
69
HFsitaAm HFAm HFAm HFAm HFAm H H C C C C C C C C C C C C T T T T T T T T T T T T T T T T T T N N N N N N N N N N i i i i i i i i i i i i i i i i i i a a a a a a a a a a a a F F L L L L L L L L L L l t F F F T T T T T L S K b G G S ShnoH S T F C D S S T T T T T o o B B S C S S S D D L G G a i i i p h a a a a a a a a a h a h h A A u a a a a a 1 s s u e G B B B h a u u u u u n S S e e o R R a a r / g b 2 2 u u u u u e 2 1 l R y S b n e m m z d 5 4 3 2 1 s g g 2 1 r V 5 4 3 a e n ? B A 5 4 3 2 1 u u sdH3 sL sS sL l a l z s d h s u e e g 1 L L F r B o s s y y a z i 1 1 y n a H a a F H2 t t o S4 S1 2 A 1 2 b g u u F 000 000 0 000 000 201 002 000 000 000 000 000 0 0 000 000 000 000 000 000 000 000 000 000 0 0 0 001 000 000 000 000 000 011 010 000 000 000 000 000 000 000 000 000 000 000 000 Grammissia 0 0 0 0 0 0 Gramicoidea 0 0 0 0 0 0 Leiopteria 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0010 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 Actinopteria Pericyclas Goniophora Spenotus 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0100 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 1000 1000 0000 0000 0000 0000 Phestia Orthonota Limoptera Glossites 0000 0000 0000 0010 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 1000 0000 0000 0000 0000 0000 0000 0000 0000 0110 0100 0100 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 Unknown Pterimorph Bellerophon Mourlonia Bembexia 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0100 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 1000 0000 0000 0000 0000 0000 0000 0000 0010 0000 0000 0001 0000 0000 0000 0000 0000 0000 0000 0000 0000 Platyceras Platystoma Cyclomena Cyrtonella 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0030 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0100 1010 0000 0000 0000 0000 0000 0000 0000 0000 1000 0000 0000 0000 0000 Paleozygoplura Holopea Glyptomaria Natochonema 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 1000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 1000 0000 0000 0000 0000 0000 0000 0000 0000 2001 1000 1001 0010 4000 0000 0010 0000 1110 0000 1000 1000 Gastropd frag Tornoceras Indet Ammonite Hyolith 00 10 01 01 00 00 01 00 00 00 00 00 00 40 00 00 00 00 00 00 02 00 00 00 00 00 00 00 00 00 00 10 00 01 00 00 00 00 00 00 00 00 02 00 00 00 00 Phacops 70 Greenops
HFsitaAm H H H H H H C C C C C C C C C C C C T T T T T T T T T T T T T T T T T T N N N N N N N N N N i i i i i i i i i i i i i i i i i i a a a a a a a a a a a a F F F F F F L L L L L L L L L L l t F F F T T T T T L S K b G G S ShnoH S T F C D S S T T T T T o o A A B B A A D L G G S C S S S D a i i i p h a a a a a a a a a h a h h A A u a a a a a 1 s s u e G B B B h a u u u u u n S S e e m m m m o R R a a r / g b 2 2 u u u u u e 2 1 l R y S b n e m m z d 5 4 3 2 1 s g g 2 1 r V 5 4 3 a e n ? B A 5 4 3 2 1 u u s s s s l a l z s d h s u e e g 1 L L F r B o L d S L s s y y a z i 1 1 y n a H H a a F H t t o S S 2 A 2 1 b g u u 4 1 F 3 2 000 000 010 000 000 000 0 000 100 000 000 000 000 000 000 000 000 000 000 000 000 200 000 000 000 000 000 000 000 000 000 0 000 000 000 000 000 000 000 000 000 000 100 000 000 000 000 Bellacartwrightia 0 0 Diplura 0 0 Indet Proetid 0000 0000 0000 1000 1000 0000 0000 0002 0000 2000 0000 0000 0020 0010 0010 6020 0000 0000 0000 0000 0000 0000 2001 0000 0000 0000 0000 0000 6000 0000 0010 1000 0110 0020 0100 0200 0210 0010 0000 0000 1000 1020 0050 0010 0000 0000 0000 Indet Trilobite Styliolinid Spyroceras Conularid 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0300 1300 0000 0000 0000 0000 0000 0001 0000 0630 0000 0000 0000 0001 0000 0000 0000 0000 0000 Auloporid Rugosa Tabulate Indet Nautiloid 00000 00000 00333 00300 00000 00130 00000 00000 00000 0063 00100 00330 00000 00000 00000 00000 06000 00000 00000 00000 00000 00000 00000 00300 00000 06000 00110 00000 00000 00600 00000 00163 00000 00000 00619 00316 00600 00600 00601 00100 00000 00600 00100 00300 00601 00100 00660 Monoplacophora Ostracod Indet Crinoid Fenestrate 1
0 Stictopora 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 1000 0000 0000 0001 0300 0000 0000 0000 0000 0000 0000 0000 0000 0003 0000 0000 Phistuliporid Taineopora Palaeschara Bryozoa frag 0200 0300 0300 0000 0300 0030 0000 0000 0000 0100 1000 0150 0000 0000 0100 0000 0100 0401 0000 0002 0700 0000 0100 0000 0100 0000 0000 002 0200 0000 0000 0262 0407 0504 0300 0300 0400 0000 0100 0007 0100 0000 0000 0007 0006 0000 0000 Unknclam Clam fragment Stroph fragments 1
0 Chonetid fragment 01 00 00 00 00 08 00 00 00 00 00 01 00 00 00 00 00 00 00 01 00 02 00 00 00 00 00 00 00 00 20 01 00 00 00 00 00 04 00 20 00 02 00 02 02 01 00 Atrypid fragment Spirifer fragment
71
B B B HFFabi H H H H H H H H H H H H H H HFShed7B HFShed7A HFShed6 HFShed5 HFShed4 HFShed3 HFShed2 HFShed1 H HFGage HFGage HFGage HFGage HFGage HFGage HFKarbd1 HFKarbd2 H H HFFaBrk HFFG8 HFFG7 HFFG6 HFFG5 HFFG4 HFFG3 HFFG2 HFFG1 o o o F F F F F i i i i i i i i i F F F F F F F F F f F F S G G C C C u b b L T o o o o o o o o o p p a a r a a a a u u u r r r r r r r r 3 s s e D n u g g 9 8 7 6 5 4 2 1 y y y S D z s t 2 1 l l l e o 1 h 3 2 1 n e 5 4 3 2 1B 1A R n d g R y 0 0 0 000 0 0 0 0 0 0 0 000 000 0 000 000 000 001 000 000 086 403 008 005 002 007 00 002 004 0 0 0 0 0 0 000 700 0 0 0 0 0 0 0 0 0 Ambocoelia 0 0 0 3 4 3 0 0 0 0 1 0 0 0 0 1 1 0 0 0 0 0 0 0 4 0 Emmanuella 1 11 8 0 0 2 2 2 0 0 1 0 9 4 0 2 0 0 0 0 2 0 0 0 0 0 0 6 Mucrospirifer 0000 0 0 0 0 0000 0000 0100 2000 0000 0000 0000 0 0 0 0000 0100 0000 0000 0000 0000 0000 0000 0000 2000 3009 0003 0008 0415 0904 0024 1005 0000 0000 0000 0000 0000 0000 0000 2000 0500 0600 0000 0000 0 3 Mediospirifer 1 2 1 1 1 3 1 1 6 2 3 8 1 100 200 3 0 Allanella 10 0 0 0 1 0 0 Cyrtina 10 0 0 0 0 0 0 Pustulatia 0 0 0 0 000 1 000 000 0 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 200 005 040 000 000 001 000 050 000 000 000 000 000 000 000 000 000 000 000 000 0 Pseudoatrypa 0 0 0 0 0 0 0 Spinatrypa 0 0 0 0 0 0 0 Elita fimbriata 0000 0000 0000 0110 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 2000 0000 0000 0000 0010 0000 0000 0000 2000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0100 Megastrophia Mesoleptostrophia Protoleptostrophia Trematospira 0610 000 000 0000 0010 0000 0008 040 000 0020 0300 0310 0500 0 0628 0230 0310 0200 0120 0300 0020 0030 0300 0 0000 0000 010 0210 0100 0020 0200 0200 0 0700 0000 0000 0000 0000 0200 0300 0820 0220 0310 0100 0600 0000 Strophodonta 1 1 1 500 340 000 Devonochonetes Longispina 1 1 1 1 1 0 1 0 3 5 Rhyssochonetes 0000 0000 0000 3000 0000 2000 0000 0000 3000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 7000 0000 0000 0000 0000 0000 0000 0000 0000 3000 7000 0000 1000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 5000 Eoschuchertella Douvillina Retichonetes Productids 0000 0000 0000 0000 0000 1 0000 0000 0000 3010 0000 0100 0700 0200 0000 0000 0200 0500 1500 0001 0000 0 0000 00 00 03 0090 00 0300 0400 0000 0500 0400 0100 00 00 00 00 00 0200 0100 0310 0100 0080 00 2500 Rhipidomella 1 1 700 720 Tropidoleptus 3 2 1 1 1 1 3 1 2 2 4 3 1 1 2 6 0 0 9 8 Leiocosta 0 0 0 0 0 0 0 0 0 0 Hypothyridina 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 0 0 0 00 0 00 00 0 0 0 0 0 0 00 00 30 00 00 00 00 00 00 00 00 00 00 00 Cupularostrum 0 0 0 0 0 0 0 0 0 0 Rhynchonellid
72
B B B HFFabi H H H H H H H H H H H H H H HFShed7B HFShed7A HFShed6 HFShed5 HFShed4 HFShed3 HFShed2 HFShed1 H HFGage HFGage HFGage HFGage HFGage HFGage HFKarbd1 HFKarbd2 H H HFFaBrk HFFG8 HFFG7 HFFG6 HFFG5 HFFG4 HFFG3 HFFG2 HFFG1 o o o F F F F F i i i i i i i i i F F F F F F F F F f F F S G G C C C u b b L T o o o o o o o o o p p a a r a a a a u u u r r r r r r r r 3 s s e D n u g g 9 8 7 6 5 4 2 1 y y y S D z s t 2 1 l l l e o 1 h 3 2 1 n e 5 4 3 2 1B 1A R n d g R y 0 2 0 000 0 0 0 0 1 0 0 0 000 000 0 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 0 1 0 0 0 0 100 000 0 0 0 0 0 0 0 0 Athyris 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Megakozlowskiella 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 "Orthis" lepidus 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0150 0000 0000 0000 1101 0000 2000 0001 0000 0000 4000 1000 0000 0000 1010 0000 0000 0000 1000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0 Lingula
0 Crania 10 Orbiculoidea
0 Lidstroemella 0 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 100 000 000 000 000 000 000 000 700 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 Pholidops
0 Unkn Inarticulata
0 Cranaena 0082 0000 0000 0000 0000 0000 0000 0010 0000 0000 0000 0000 0000 0062 0060 0000 0051 0060 0062 0000 0061 0000 0041 0022 0000 0021 0023 0000 0000 0030 0000 0000 0000 0020 0000 0040 0030 0000 0091 00 0000 0000 0031 0000 0030 0030 Cryptonella Unkn Terebratulid 1
3 Nuculoidea
0 Nuculites 0000 0000 0000 0000 0000 0000 0000 0003 0000 0000 0002 0000 0002 0001 0300 0000 0007 0002 0007 0001 0000 0000 0001 0102 0000 0000 0010 0003 0000 0000 0010 0000 0000 0005 0003 0006 000 0005 0005 0004 0000 0000 0000 0004 0000 0006 Nuclea Modiomorpha Modiella 1
1 P. constricta 1000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 1000 0000 0000 0000 3000 0000 2000 3000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 2000 0000 0000 0000 0000 0000 0000 0000 0000 0000 1000 P. emarginata P. filosa Pholidella Cimitaria 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 1000 0000 0300 0000 0000 0000 0000 0000 0000 0000 1000 0000 1000 1000 0000 0000 0000 0000 0000 0000 0500 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 Tellinopsis Cypricardella Phthonia Meristella 00 00 00 00 00 00 00 00 00 02 00 00 00 00 00 00 00 00 00 00 00 00 01 00 00 00 00 00 00 00 00 00 00 00 00 02 00 00 00 00 00 00 00 01 01 00 Mytlops Ontaria
73
B B B HFFabi H H H H H H H H H H H H H H HFShed7B HFShed7A HFShed6 HFShed5 HFShed4 HFShed3 HFShed2 HFShed1 H HFGage5 HFGage4 HFGage3 HFGage2 HFGage1B HFGage1A HFKarbd1 HFKarbd2 H H HFFaBrk HFFG8 HFFG7 HFFG6 HFFG5 HFFG4 HFFG3 HFFG2 HFFG1 o o o F F F F F i i i i i i i i i F F F F F F F F F f F F S G G C C C u b b L T o o o o o o o o o p p a a r a a a a u u u r r r r r r r r 3 s s e D n u g g 9 8 7 6 5 4 2 1 y y y S D z s t 2 1 l l l e o 1 h 3 2 1 n e R n d g R y 1 0 0 0 0 0 0 0 0 0 0 0 0 0 000 000 000 0 000 000 000 000 000 000 000 000 000 000 010 000 000 000 000 0 0 0 0 0 0 0 0 0 0 000 000 0 Grammissia 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Gramicoidea 0 0 0 0 0 1 0 0 1 1 0 0 0 0 0 0 1 3 2 0 0 0 0 0 0 0 Leiopteria 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 1000 1000 2100 2000 2000 1000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 Actinopteria Pericyclas Goniophora Spenotus 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 2000 0000 0000 3000 2000 4000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 Phestia Orthonota Limoptera Glossites 0000 0000 0000 0000 0000 0000 0000 0100 0000 0000 0000 0000 0010 0000 0010 0000 0000 0000 0000 0000 0010 0010 0010 0000 0000 1000 0000 0000 0000 0000 0000 1010 0100 0100 0300 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 Unknown Pterimorph Bellerophon Mourlonia Bembexia 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 Platyceras Platystoma Cyclomena Cyrtonella 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0100 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 Paleozygoplura Holopea Glyptomaria Natochonema 3000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 1000 0000 0000 0000 0120 0010 1000 0000 0000 0000 0000 0000 0000 0000 1000 0000 0000 0000 0010 0000 0020 0000 0000 0000 0000 0000 0000 0000 0000 Gastropd frag Tornoceras Indet Ammonite Hyolith 01 00 00 00 00 00 00 00 00 00 01 11 00 10 10 02 00 02 00 00 00 00 00 00 00 00 01 00 00 00 00 00 00 00 00 01 01 02 02 11 00 00 10 00 01 10 Phacops 74 Greenops
B B B H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H o o o F F F F F F i i i i i i i i i F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F f F F S G G F C C C u b S S S S S S S S b G G G G G G K K L T F F F F F F F F F o o o o o o o o o p p a a h h h h a G G G G G G G G a h h h h r a a a a u u u a a a a a a a a r r r r r r r r 3 s s e b B D n u e e e e e e e e g g 9 8 7 6 5 4 2 1 g g g g g g r r y y y 8 7 6 5 4 3 2 1 D S z b b i d d d d d d d d s t 2 1 e e e e e e l l l r e o 1 h 3 2 1 n d d k 7 7 6 5 4 3 2 1 e 5 4 3 2 1 1 R n d g R 1 2 B A B A y 000 000 000 000 000 000 000 000 000 000 000 000 000 100 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 Bellacartwrightia Diplura Indet Proetid 0010 0000 2000 0120 0000 0000 0000 0000 0000 0000 0010 0000 0000 0000 0000 0000 1000 0000 0000 0000 0200 0030 0000 0000 0000 0000 0020 0000 0000 0000 0000 0000 0010 0000 0000 0000 0000 0000 0010 0000 0000 0000 1020 0300 2000 0000 Indet Trilobite Styliolinid Spyroceras Conularid 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 Auloporid Rugosa Tabulate Indet Nautiloid 00000 00000 00000 00000 00000 00000 06000 00000 00033 00000 00000 00000 00000 00000 00000 00000 00611 00110 00000 00000 00000 00000 00000 00000 00000 00000 00310 00000 00000 00000 00000 00000 00000 00000 00000 00000 00000 00000 00010 00000 00100 00000 00000 00000 00000 00220 Monoplacophora Ostracod Indet Crinoid Fenestrate Stictopora 0030 0030 0030 0030 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 Phistuliporid Taineopora Palaeschara Bryozoa frag 0300 0000 0300 0300 0000 0300 0000 0000 0000 0000 0000 0000 0000 0000 0000 0300 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0303 0300 0300 0000 0000 0000 0000 0000 0000 0000 0003 0000 0000 0000 0000 0000 0000 0000 0000 Unknclam Clam fragment Stroph fragments Chonetid fragment 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 03 00 00 00 00 00 00 00 00 00 00 00 00 00 03 00 00 03 00 00 00 06 00 06 00 00 00 00 00 00 00 Atrypid fragment Spirifer fragment 75
b b N G G b b G G W D N X H D J C S M W W G G G G F F F K K K K K B B B B B B B B B B B B u a a a a h a a a a a a a a a a o o o o o o w a a a a e E R o e r r r r a a a a a i H B B B T L T T B B B o o o o e R F F F F F F s s s s s g g g g R R n T T T T T S G S d L t G S S L G K S S S u a u u S S S i e 3 2 1 e r r r i i i i i i p o D a a a a a b t h t F l s s s s s s L G F N s N S S p p p p p G o W a W P S w w w H N u u u u u e a a a S / 4 3 3 2 1 H H N / H B n a e e e e e D D S T z g a g B o w S 5 4 3 2 1 E r h B A a a a i e V B n i z z z z z e i s 2 E e g F h i f e e a h R R l l l F e F L d N L 2 1 3 2 1 F r s S n t t t d T S o N n o o i 2 1 s o L h s L 000 100 000 000 000 000 000 000 000 011 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 022 700 000 000 300 000 000 500 000 300 000 000 000 000 000 000 400 Athyris Megakozlowskiella "Orthis" lepidus 0000 0000 2000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0010 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 5000 0000 0000 0000 0000 0000 0000 2000 0000 0010 Lingula Crania Orbiculoidea Lidstroemella 000 000 000 000 000 000 010 000 000 000 000 010 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 010 000 000 000 000 000 000 000 000 000 000 000 000 000 000 100 Pholidops Unkn Inarticulata Cranaena 0000 0010 0010 0000 0000 0010 0000 0030 0000 0000 0010 0030 0020 0000 0001 0010 0000 0030 0011 0010 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0020 0000 0000 0042 0001 1001 0000 0000 0010 0000 0000 0000 0000 0000 0061 Cryptonella Unkn Terebratulid Nuculoidea Nuculites 0000 0001 0003 0004 0000 0000 0002 0000 0000 0000 0006 0000 0000 0000 0000 0101 0000 0000 0000 0000 0100 0000 0002 0000 0000 0002 0001 0001 0000 0000 0000 0000 0000 0006 0001 0000 0000 0000 0000 0000 0000 0000 0003 0003 0001 Nuclea Modiomorpha Modiella P. constricta 0000 0000 0000 0000 0000 0000 0000 0000 0000 0010 0000 0000 0000 1000 0000 1000 0000 0000 0000 2000 2100 0000 0000 0000 0000 0000 1000 0000 0000 0000 0000 0000 0000 0000 2000 3200 0100 3000 1000 0000 1000 0000 1100 0000 0001 P. emarginata P. filosa Pholidella Cimitaria 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0001 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 1000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0100 0000 0000 Tellinopsis Cypricardella Phthonia Meristella 00 04 00 00 00 00 00 01 02 00 00 00 00 00 00 00 10 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 Mytlops Ontaria 76
b b b N G G b b G G W D N X H D J C S M W W G G G G F F F K K K K K B B B B B B B B B B B B u a a a a a h a a a a a a a a a a o o o o o o w a a a a e E R o e r r r r a a a a a i H B B B T T L T T B B B o o o o e R F F F F F F g g g g R R n s s s s s T T T T T S G S d L t G S S L G K S S S u u a u u S S S i e 3 2 1 e r r r i i i i i i p D o a a a a a b t h t F l s s s s s s L G F N s H N S S p p p p p G o W a W P S w w w H N u u u u u e a a a S H / 4 3 3 2 1 H N / H B n a e e e e e S D D T z g g B a o w S 5 4 3 2 1 o E r h B A a a a i e V B n i z z z z z e i s 2 E e g F h i e e f a h G l l l F R R e L F d N L F 2 1 3 2 1 r s S n t t t d T S o N n o o i l 2 1 s o L h s L 000 000 000 000 000 000 000 000 000 000 000 000 000 200 000 000 000 000 011 200 000 000 000 000 000 000 000 000 000 000 000 100 000 000 000 003 000 000 000 000 000 100 000 000 000 000 Bellacartwrightia Diplura Indet Proetid 1 1 1 0000 0100 1000 0000 2301 0000 2110 0000 0000 0000 0000 1010 0000 5300 0001 1600 2000 7010 1000 0000 3000 0000 0020 0010 0000 0000 1000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 1000 0000 0000 0040 0020 0000 0000 0000 1020 Indet Trilobite Styliolinid Spyroceras Conularid 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0600 0000 0000 0 0000 0000 0200 0000 0000 2800 0300 0000 0000 0000 0000 0000 0000 0 0000 0000 0000 2 0000 0000 0000 0000 0301 0000 0000 0000 0200 0000 0000 0000 0000 0000 Auloporid 1 1 1 002 000 400 Rugosa Tabulate Indet Nautiloid 00100 01000 00000 00000 00000 10000 00000 00111 00000 00000 00660 00300 00000 00121 00613 00130 00000 00000 00000 00000 00000 00000 00000 06000 00672 00000 00000 00683 00000 00000 00000 00660 00360 00000 00000 00000 00000 00000 00000 00333 00010 00630 00665 00660 00360 00000 Monoplacophora Ostracod Indet Crinoid Fenestrate Stictopora 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 3000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 2000 0000 0000 0000 0000 3000 0000 0000 0000 0000 0000 1000 0000 0000 0000 0000 0000 0000 0000 0000 Phistuliporid Taineopora Palaeschara Bryozoa frag 0000 0000 0000 0000 0000 0000 0000 1000 0000 0300 0100 0000 0000 0000 3000 0000 0000 0100 0100 0000 0000 0000 0000 0000 0170 0300 0000 0200 0000 0000 0000 0000 0200 0000 0000 0000 0000 0000 0000 0000 0000 0100 0000 0300 0100 0000 Unknclam Clam fragment Stroph fragments Chonetid fragment 00 00 00 00 00 00 02 00 00 03 00 00 00 00 00 00 00 00 00 00 00 00 00 00 04 00 00 01 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 02 00 00 Atrypid fragment Spirifer fragment
77
b b b N G G b b G G W D N X H D J C S M W W G G G G F F F K K K K K B B B B B B B B B B B B u a a a a a h a a a a a a a a a a o o o o o o w a a a a e E R o e r r r r a a a a a i H B B B T T L T T B B B o o o o e R F F F F F F s s s s s g g g g R R n T T T T T S G S d L t G S S L G K S S S u u a u u S S S i e 3 2 1 e r r r i i i i i i p o D a a a a a b t h t F l s s s s s s L G F N s H N S S p p p p p G o W a W P S w w w H N u u u u u e a a a S / 4 3 3 2 1 H H N / H B n a e e e e e D D S T z g a g B o w S 5 4 3 2 1 o E r h B A a a a i e V B n i z z z z z e i s 2 E e g F h i f e e a h G l R R l l F e F L d N L 2 1 3 2 1 F r s S n t t t d T S o N n o o i l 2 1 s o L h s L 000 000 000 000 000 000 000 000 000 000 000 000 200 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 003 000 000 Grammissia Gramicoidea Leiopteria 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0010 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0001 0000 0000 0000 0000 0000 0000 0000 0000 Actinopteria Pericyclas Goniophora Spenotus 0000 0000 0000 0000 0010 0000 0000 0001 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0010 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 Phestia Orthonota Limoptera Glossites 0000 0000 0000 0000 0000 0120 0000 0001 0000 0000 0000 0000 0000 0000 0000 0000 0000 1000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0100 0100 0000 0000 3000 0000 0000 0000 0000 1000 0000 0000 0000 0000 0000 0000 0000 0000 0000 Unknown Pterimorph Bellerophon Mourlonia Bembexia 0000 0000 0000 0000 0000 0000 0000 0000 0100 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0002 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 Platyceras Platystoma Cyclomena Cyrtonella 0000 0000 0000 0000 0000 0000 0000 0000 1000 0000 3000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0002 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 9000 0000 0001 0000 0000 0000 0000 0010 1000 Paleozygoplura Holopea Glyptomaria Natochonema 0000 0000 0000 0000 0200 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 1000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0100 0100 0000 Gastropd frag Tornoceras Indet Ammonite Hyolith 00 00 00 00 00 00 00 00 00 00 00 01 00 00 00 00 00 00 00 00 02 00 00 00 00 00 00 00 00 10 00 90 00 00 00 00 00 01 00 00 01 00 00 00 01 00 Phacops 78 Greenops
s b u S S S b H D D D D b s b u S S S b H D D D D b u u a p a a a p a a h h h h h h i e e e e i e e e e b b T N D T N D F F N N e e e e e e R S R R R S R R D D d d d d d d o o u u e e e e S S e e y y y y y y W J S W J S J J S S L N L N R R L L h h S J S S J S u u u u F F P P u u s s e e s s S S W W k D k D n n e e n n N N a a N i N i n d n d S S a a r r e e t t e e b b e e n n e s e s D D L L n n L L d d r r R R 000 0 000 000 000 000 000 006 0 000 002 00 000 300 000 000 000 000 000 000 000 0 000 000 Athyris 002 0 Ambocoelia 1 1 1 1 64 41 91 Megakozlowskiella 1 Emanuella 1 9 "Orthis" lepidus 4 Mucrospirifer 0000 0000 0000 0000 0000 0004 2200 0000 0000 000 0000 0000 0200 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 Lingula 0030 Mediospirifer Crania Allanella Orbiculoidea Cyrtina 2
Lidstroemella 4 Pustulatia 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 Pholidops 002 Pseudoatrypa Unkn Inarticulata Spinatrypa Cranaena Elita fimbriata 0000 0000 0000 0000 0000 0000 0000 0000 0070 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 Cryptonella Megastrophia Unkn Terebratulid Mesoleptostrophia Nuculoidea Protoleptostrophia Nuculites Trematospira 0000 0001 0000 0000 000 0000 0000 000 0005 0000 0000 0000 0000 0004 0000 000 000 100 0000 0000 0000 0000 0000 0000 0201 0000 Nuclea Strophodonta Modiomorpha Devonochonetes Modiella Longispina 1 1 1 1 1 5 1 P. constricta 1 4 4 Rhyssochonetes 0000 0000 0000 0000 0000 0000 0000 2000 2000 0000 0000 0000 0000 0000 0000 1000 3000 0000 0000 0000 0000 0010 0000 7308 4000 0000 P. emarginata Eoschuchertella P. filosa Douvillina Pholidella Retichonetes Cimitaria Productids 0000 0000 0000 0800 0000 0000 0000 0200 0100 0000 0000 00 0000 0000 0000 0000 0100 0090 0000 0000 0000 0000 0020 0000 0000 0000 Tellinopsis Rhipidomella Cypricardella Tropidoleptus 2
Phthonia 1 Eumetabolotoechia
Meristella 0 Hypothyridina 00 00 00 00 00 00 00 0 00 00 00 00 00 00 00 00 00 00 00 00 00 00 00 40 00 00 Mytlops Cupularostrum
Ontaria 0 Rhynchonellid
79
s b u S S S b H D D D D b s b u S S S b H D D D D b u u a p a a a p a a h h h h h h i e e e e i e e e e b b N D T T N D F F N N e e e e e e R R R S R R R S D D d d d d d d o o u u e e e e S S e e y y y y y y W J S W J S J J S S L N L N R R L L h h S J S J S S u u u u F F P P u u s s e e s s S S W W k D k D n n e e n n N N a a N i N i n d n d S S a a r r e e t e t e b b e e n n e s e s D D L L n n L L d d r r R R 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 000 Bellacartwrightia 000 Grammissia Diplura Gramicoidea Indet Proetid Leiopteria 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 3000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 6000 0000 Indet Trilobite 0000 Actinopteria Styliolinid Pericyclas Spyroceras Goniophora Conularid Spenotus 0000 0000 0000 0000 7000 0000 0000 0000 0000 0000 0000 5000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 Auloporid Phestia Rugosa Orthonota Tabulate Limoptera Indet Nautiloid Glossites 0000 00000 0000 0410 0000 0000 00000 0000 0000 00000 00000 00000 0000 0000 0000 00000 00010 0000 00000 00000 00000 0000 00300 0000 00330 00000 Monoplacophora Unknown Pterimorph Ostracod Bellerophon Indet Crinoid Mourlonia Fenestrate Bembexia 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 Stictopora 0000 Platyceras 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 Phistuliporid Platystoma Taineopora Cyclomena Palaeschara Cyrtonella 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 Bryozoa frag 0000 Paleozygoplura 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 Unknclam Holopea Clam fragment Glyptomaria Stroph fragments Natochonema 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 Chonetid fragment 0000 Gastropd frag 00 00 00 00 00 00 00 00 00 00 00 00 00 Atrypid fragment Tornoceras Spirifer fragment Indet Ammonite
Hyolith 00 00 00 00 00 00 00 00 00 00 00 00 00 Phacops 80 Greenops