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University of Nevada, Reno

Paleolimnology and of the Miocene Quincy Diatomite Deposit

A thesis submitted in partial fulfillment of the requirements for the degree of Master of Science in Geology

By Anthony Joseph Menicucci

Dr. Paula J. Noble / Thesis Advisor

August 2010

THE GRADUATE SCHOOL

We recommend that the thesis prepared under our supervision by

ANTHONY JOSEPH MENICUCCI

entitled

Paleolimnology and Paleontology of the Miocene Quincy Diatomite Deposit

be accepted in partial fulfillment of the requirements for the degree of

MASTER OF SCIENCE

Dr. Paula J Noble, Advisor

Dr. Alan Wallace, Committee Member

Dr. Mary Peacock, Graduate School Representative

Marsha H. Read, Ph. D., Associate Dean, Graduate School i

Abstract

The Quincy Diatomite deposit, central Washington, USA, is a middle Miocene

(~15 Ma) freshwater lacustrine deposit located between flows of the Columbia River

Basalt Group. Three localities along the western margin of the deposit are examined and the flora are described at the species level, with 84 species recognized. Of these species, one is new, Fragilariforma intortus, and one is elevated in rank to the species level, Tetracyclus williamsensii. One species, Pseudostaurosira brevistriata var. subcapitata, is also is transferred to a new genus for taxonomic clarification. Although rare, three species of the genus Sellaphora occur in the Quincy diatomite deposit, making this occurrence the oldest for the genus in the record. The northwestern United

States endemic fossil species Ellerbeckia baileyi is also observed in the deposit, and is abundant in particular stratigraphic intervals.

Stratigraphic and geographic variation within the deposit is also examined. The three lithologic units recognized in the deposit, the Four Crude, Bottom Crude, and Top

Crude, are distinguished by subtle lithologic differences and contain distinct diatom floras. Textural properties are largely a function of floral differences between units.

Diatom assemblages also vary geographically along the western margin of the basin particularly between localities of Top Crude in northern outcrops of the Ancient

Park versus more southerly localities in the Frenchman Hills and Gorge Amphitheater.

Distinctions between these assemblages are confirmed through analysis of point count data using nonMetric Multidimensional Scaling and cluster analyses.

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Some species within the flora, primarily Aulacoseira granulata and

Staurosira construens var. venter, are used as ecologic proxies to interpret the paleolimnology of the Ancient Quincy as an alkaline, eutrophic to hypereutrophic lake with high Total (TP), and sufficient dissolved Silica (Si) levels capable of sustaining a large population.

Three stages of development in lake ecology, directly influenced by the regional geology and geography, are recognized. The earliest stage, represented by the Four

Crude, was deposited in a series of small-interconnected pocket lakes that were monomictic, eutrophic, slightly alkaline, and had a depth of greater than 15 meters. This lake stage was succeeded by Bottom Crude deposition and is interpreted to represent the period of maximum shallowing, creating a polymictic lake of less than 10 meters depth and dominated by benthic species. The latest stage of the lake, represented by the Top

Crude, represents the deepest and broadest expansion of the lake and shows variable development of lake stratification through its deposition. Tectonic uplift of the western

Frenchman Hills Anticline combined with overall subsidence of the Quincy Basin during intermediate and late stages of lake development is interpreted to be the driver for changes in lake bathymetry.

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Acknowledgments

This work would not have been possible if it were not for the assistance and support of several people, I owe a great debt to all who have helped me in this endeavor.

I’d like to thank the Nevada Petroleum Society for their grant awards for the 2007 and

2008 years and the Paleontological Society for their 2007 grant. I would also like to thank my advisor, Paula Noble, for her help and assistance during this undertaking; she taught me to write like a scientist, and had endless patience for my ambition, helping me to focus my efforts and develop this work. My committee, Alan Wallace and Mary

Peacock area also due a great deal of thanks for their consultations as I progressed through the project. Many thanks go to World Minerals inc. and Mike Houseman and the

United States Army Corps of Engineers for access to materials and all their assistance both in the field and the lab; this work would not have been possible without their help.

I’d also like to thank my family for all their support and encouragement through the entire process. Many thanks go to my wife Nichole, for her support and willingness to put up with me through this undertaking, and her support as I progress in my professional and academic careers. Finally, all the people at Iowa Lakeside Laboratory, including

Sarah Spaulding, Mark Edlund, and Steve Juggins for their consultation and teaching efforts; thank you all.

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Table of Contents

Abstract i

Acknowledgements iii

Table of Contents iv

List of Tables vii

List of Figures viii

Chapter 1: Paleolimnology and Paleontology of the Quincy Diatomite

I. Introduction 1

II. Regional Geology 6

III. Methods 16

IV. Observations 21

V. Limnologic Interpretations 40

VI. Summary and Conclusions 58

Chapter 2: Systematics and Species Descriptions with rare species list

I. Systematics (by Genus)

a. Aulacoseira 62

b. Melosira 66

c. Actinocyclus 67

d. Ellerbeckia 70

e. Diatoma 72

f. 75

g. Fragilariforma 78

h. Staurosirella 80

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i. Meridion 83

j. Pseudostaurosira 84

k. Staurosira 86

l. Synedra 88

m. Tetracyclus 89

n. Eunotia 96

o. Cavinula 99

p. Neidium 100

q. Sellaphora 101

r. 102

s. Diploneis 104

t. 105

u. Cymbella 107

v. Placoneis 109

w. Gomphonema 112

x. Gomphosphenia 117

y. Planothidium 118

z. Nitzschia 122

aa. Extremely Rare Species List 124

II. References 126

III. Plates #1-23 140

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APENDICES

a. Appendix 1– Sample numbers and Sampled Cores 186

b. Appendix 2– Specimen Picture X-Y coordinates 191

c. Appendix 3– Point Count Data 197

d. Appendix 4–Error Calculation Counts 206

e. Appendix 5– Measurements of Width for Grain Size Calculation 217

f. Appendix 6– Extended Methodologies 227

g. Appendix 7 – Organic Carbon Test Results 236

h. Extended Bibliography – Includes References used but not cited 238

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LIST OF TABLES

1. Table 1: UTM Coordinates for all sample sites 17

2. Table 2: Species Occurrence list/Key to species for NMDS 35

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LIST OF FIGURES

1. Figure 1: Locality Map of the Quincy Diatomite deposit 4

2. Figure 2: Stratigraphy of the Columbia River Basalts 9

3. Figure 3: Stratigraphy of the Quincy Diatomite deposit 11

4. Figure 4: Relative Abundance Stratigraphic Plot: Frenchman Hills 23

5: Figure 5: Relative Abundance Stratigraphic Plot: Ancient Lakes Park 29

6: Figure 6: NMDS Plot of Species and Samples 33

7: Figure 7: Cluster Analysis – Wards Method of Samples 39

8: Figure 8: Modeled Extent of Quincy Ancient Lake 55

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Chapter 1: Paleolimnology and Paleontology of the Quincy Diatomite deposit

I. Introduction

The Quincy Diatomite deposit is a Miocene lacustrine deposit that occurs within the Columbia River Basalt Group in central Washington. This deposit is located in the

Quincy Basin in the northern part of the Yakima Fold Belt (Figure 1). The diatomite is industrial grade and hyper pure (> 90% pure diatom material), and it has been actively mined for the last 50 years in several localities, primarily in and around the Frenchman

Hills area.

Although some work has been published on the Quincy flora, no one has comprehensively studied the species composition and floral variation either stratigraphically or geographically. Vanlandingham (1964, 1967) studied Miocene from the central Washington area including some small outcrops to the south of the Frenchman Hills (Figure 1A). His study broadly detailed the floral assemblages of the entire central Washington region through multiple time periods in the Miocene, but specific characterization of the Quincy flora and interpretation of its ecology was left for future research. Kociolek and Spaulding (2002) examined material from the northern areas of the deposit (Figure 1A) in their paper where they suggested the possibility of a single species with multiple morphologies expressed as Ellerbeckia sp. and Actinocyclus sp.. However, their study was topical in nature and the entire flora was not described.

Houseman (2006) reported some preliminary data on relative abundance counts, primarily at the genus level. His findings showed stratigraphic variation at the genus level, and illustrated fundamental differences between each of the three units within the

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Quincy Diatomite deposit. Houseman’s findings indicated further investigation of the Quincy flora should provide useful paleolimnological data.

The Quincy Diatomite deposit holds particular significance because the extremely good diatom preservation provides great potential to aid in understanding geologic and paleolimnologic influences in a Miocene diatomite-bearing basin. Such basins exist commonly in the Western US (Lohman, 1957, Bradbury and Krebs, 1995, Bolm et al.,

2003), yet little work has been done to characterize whole assemblages at the species- level and then use any observed changes in paleolimnological reconstructions of diatomites. Further, the Quincy deposit is a substantial sedimentary deposit located between flows of the Columbia River Basalt Group, and as such is a window into middle

Miocene during the eruptive phase of this thick sequence of basalt flows.

This study presents the findings from three localities along the western margin of the Quincy Basin that characterize species level changes between sampling sites. These data are used to interpret the paleolimnology of the Quincy Diatomite (including the paleolake nutrient budget, lake and basin evolution, and the relationship of the deposit to the regional geologic history) based on the fossil floral assemblages, and evaluate the robustness of the interpretations in the absence of the original lacustrine body.

Interpretations of the broad-scale paleolimnology are made using known ecologic preferences of several extant species, and an overall model of basin development is presented. Inferences are also made on the paleoecology of a number of extinct genera, based on their associations with other diatom species, including Ellerbeckia baileyi

Crawford and Sims, a diatom reported from other Miocene deposits in the Pacific

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Northwest and which occurs in this diatomite in great abundance at specific stratigraphic intervals.

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Figure 1: Locality map of the Quincy diatomite deposit. 1A shows the location of the modern and Miocene Quincy basins (with the modern Quincy basin boundary defined by the dashed line), showing sample localities and major and geographic structural features discussed in text. Moses Lake and Potholes Reservoir are outlined in blue on the Eastern side. Figure 1B is an enlarged view of the Frenchman Hills, showing World Minerals Inc. core locations and the presence or absence of the lowest two units for those cores. Stars indicate the locations of the cores sampled for this study. Figure 1B, shows cores taken in the Frenchman Hills where the basal Four Crude or the younger Bottom Crude were encountered; closed dots encountered both the Four Crude and Bottom Crude, open dots encountered the Four Crude only. The upper Top Crude is present in all locations sampled on the anticline flanks. Gray boxes in both figures are sample locations for Kociolek and Spaulding (2002); gray ovals in Figure 1A are sample locations for Vanlandingham (1964). FH = Frenchman Hills, GA = Gorge Amphitheater, ALP = Ancient Lakes Park, ML = Moses Lake.

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II. Regional Geology Regional Tectonics

The Quincy Basin geology is strongly influenced by the regional structural features. The Quincy Basin is known to have subsided at a pace consistent with the erupting volume of the Columbia River Basalt flows (Reidel, 1984, Martin, 1991). This subsidence permitted a continual releveling of the topography by the Columbia River

Basalt Group (CRBG) during the eruptive phase (Reidel, 1984, Martin, 1989, Martin

1991), and provided a catchment for in between basalt emplacement in the

Yakima Fold Belt (YFB). This subsidence has continued from initiation of CRBG eruption on the Columbia Plateau through present times (Reidel et al. 1989, Hooper

1997), as demonstrated by thinned basalts atop the anticlinal ridges, and the facultative damming of some CRB flows against these anticline ridge tops (Reidel 1984, Reidel et al

1989, Watters 1989), thus proving a direct overlap in space and time.

The Yakima Fold Belt subprovince is one of four structural subprovinces on the

Columbia Plateau, encompassing the western-central area. It is just west of the Pasco

Basin (where the CRBg sequence is thickest) and incorporates the Quincy Basin on its eastern margins (Campbel 1989, Reidel et al. 1989, Watters 1989, Hooper 1997). These anticlinal folds are associated with emergent thrust faults in the back limbs and are deformed by the through-going, N-NW trending, right-lateral structure of the Olympic-

Wallowa lineament (OWL) (Reidel 1984, Hooper and Conrey 1989, Reidel et al. 1989,

Watters 1989). Folds of the YFB are also observed to have very regular spacing through

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most of the subprovince, particularly in the northern areas of the Saddle Mountains and Frenchman Hills (Reidel 1984, Watters 1989).

Even though the area of the YFB was generally undergoing subsidence, the anticlinal folds were still active and continued growth through eruption of the CRBs (as seen, again, by the thinned basalts atop the anticlines), with each successive flow releveling the topography of the YFB subprovince. This resulted in a regional subsidence with small amounts of local shortening (Reidel 1984, Reidel et al. 1989).

Basalt Stratigraphy The Columbia River Basalt Group is composed of basaltic lavas extruded in the

Miocene between 17.0 and 6.0 Ma (Hooper and Riedel, 1989, Tolan et al., 1989, Hooper,

1997). Within this group is the Wanapum Formation, which accounts for approximately six percent of the total volume of the CRBG (Tolan et al., 1989). The Wanapum

Formation is divided into five members, including, in ascending stratigraphic age, the

Lookingglass, the Frenchman Springs (15.3-15.5 Ma), the Shumaker Creek, the Roza, and the Priest Rapids members (14.5 Ma) (Figure 2) (Hooper, 1997).

The Quincy Diatomite deposit occurs between the underlying Roza and overlying

Priest Rapids Members of the Wanapum Basalt (Figure 3). Isotopic dates on the

Frenchman Springs and the Priest Rapids members generally constrain the depositional age of the Quincy Diatomite deposit between 15.3 and 14.5 Ma. However, the Roza

Member, which overlies the Frenchman Springs and directly underlies the diatomite, has not been isotopically dated, so the Quincy Diatomite deposit is constrained only by

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published dates on the basalt stratigraphy (Hooper 1997) and the actual period was undoubtedly shorter.

The Roza Member has several named flows (Martin, 1989, Martin, 1991). The

Roza IIa flow universally underlies the diatomite both in outcrop and where the base of the diatomite is encountered during drilling. Most commonly the Priest Rapids Member overlies the diatomite (Groiler and Bingham, 1971, Tolan et al., 1989), with the Roza IIb flow locally overlying the diatomite at the Gorge Amphitheater. The Priest Rapids

Member thermally altered the top of the diatomite in the east and created a baked paleosol cap on the diatomite in the Frenchman Hills. The Roza IIb basalt created a cap of palagonitized basalt at the Gorge Amphitheater, with this palagonatized zone also incorporating small amounts of the diatomite at the basalt-deposit interface.

Finally, evidence near the Gorge Amphitheater shows the Roza IIb basalt flow interacted with wet of the ancient Quincy Lake via extensive palagonite development at the Gorge Amphitheater (Figure 3). Houseman (2006) thought that this flow would have leveled the topography of the region and effectively ended the ancient

Quincy Lake. However, the Roza IIb flow is very restricted in its distribution and is absent throughout most of the Quincy Basin (Martin 1989, 1991). This limited geographic distribution makes it more difficult to conclude a complete releveling of the topography by the Roza IIb; most exposures of the diatomite are overlain by Priest

Rapids basalt, with no evidence of Roza IIb (and the accompanying palagonite) in the

Frenchman Hills or the Quincy Ancient Lakes Park (Figure 3).

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Figure 2:. Stratigraphy of the Columbia River Basalt Group, including magnetic orientation and radiometric dates of the units. The Quincy diatomite deposit occurs between the Priest Rapids and Roza members of the Wanapum Formation. Taken from Hooper (1997).

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The Quincy Diatomite

The Quincy Diatomite deposit itself is large, industrial grade, and formed in a freshwater environment. It is commercially mined because it is so pure and has no siliciclastic interbeds (i.e. no uneconomic sedimentary intervals), with minimal lenses of diagenetic chert observed throughout the section. Stratigraphically, the Quincy Diatomite deposit can be subdivided into three ascending units, the Four Crude, Bottom Crude and

Top Crude (Figure 3). These units were defined by geologists at World Minerals

(Houseman 2006) and are distinguished in the field by slight color changes in outcrops, mine pit walls, or drill cores. Subtle textural differences that can be felt when crushing the diatomite between the fingers are also used to distinguish each unit. A distinct diatom assemblage corresponds to different levels of coarseness to the touch within each unit, and each unit is described below. Only one marker bed, a mafic air-fall tuff that is locally called the “Green Sand” is present near the base of the Top Crude unit (Figure 3).

The diatomite is white to slightly tan, highly porous, poorly indurated in both core and outcrop, and it contains small and variable amounts (1-2%) of organic carbon (Appendix

7). The deposit is 3-12 meters thick along the western margin and thins to less than 1 meter thick along the eastern margin at locations near Moses Lake, approximately 55 kilometers away. The northern boundary is just south of the Beezely Hills and the town of Quincy, and the southern margin appears to be near the base of the Saddle Mountains in the Royal Slope. Overall, the deposit covers an area over 2600 kilometers2 (Figure

1A) and based on known outcrop locations and occurrences in drill cores, is distributed across the entire Miocene Quincy basin.

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Figure 3: Stratigraphy of the Quincy Diatomite deposit at the three western localities that were sampled for this study. The pink bar in the Frenchman Hills shows the baked paleosol cap at this locality. The Gorge Amphitheater section shows the palagonite breccia cap from the Roza IIb basalt in the green polygons. The green line in the Top Crude shows the location of the “Green Sand” mafic tuff. The stratigraphic position and sample numbers for samples collected and examined for this study are shown on the left side of each locality. Modified from Houseman (2006).

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The Four Crude and Bottom Crude units are restricted in their distribution, and they are present along a northwesterly trend only in the western parts of the

Frenchman Hills (Houseman, 2006). These two units are rarely visible in outcrop. Drill cores show the thickness of the both Four Crude and Bottom Crude varies from zero to two meters between sections. The distribution of these two units is patchy between the sampled drill core localities, with some cores showing thickened or thinned Four Crude and only pockets of the Bottom Crude. Plots of unit thickness from drill core logs suggest these lower units most likely formed in small depressions that are approximately

1-2 km2 in size, and often spaced 1-2 kilometers apart (Figure 1B).

The diatomite shows a clear northwesterly orientation in the Frenchman Hills.

Evidence for a northwest trending fabric comes from unpublished drill core data from

World Minerals Inc. that shows the Four Crude and Bottom Crude occur along a northwest trend in the Frenchman Hills (Houseman, 2006) (Figure 1b). This trend is visible in sediments underlying the Columbia River Basalts (Campbell, 1989), and it has been mapped in some of the overlying basalts (Hooper, 1997) and as an aeromagnetic anomaly (Reidel, 1984). This trend, however, is much more subtle than the dominant east – trending Yakima Fold Belt in the area (Reidel, 1984, Watters, 1989). The Quincy

Diatomite deposit also thickens to the west, which is consistent with a northwest- trending, down-slope direction followed by Wanapum Basalt flows as they progressively filled the Quincy Basin (Schminke, 1967, Hooper, 1997).

The Top Crude is the most widespread unit in the Quincy Basin. It is encountered along most of the east-west trend of the Frenchman Hills, at the Gorge Amphitheater, and

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at the Quincy Ancient Lakes Park. The Top Crude varies in thickness (0.5 – 4 meters), although a minimum thickness of 2.5 meters appears to be consistent along the western margin of the Quincy Basin near the Columbia River. Gravity flow deposits occur as ~2-3mm thick layers containing rip-up clasts of diatomite in a diatomite matrix, and are subtle features visible during careful examination throughout the Top Crude in the Frenchman Hills. These gravity deposits are not observed in northern sampling localities. The Top Crude also displays a paleosol in the Frenchman Hills. This paleosol appears to be baked, and exhibits features such as incorporation of basal diatomite material, and amorphous silica fragments. Outcrops containing the paleosol show an overall terra-cotta color underlying the Priest Rapids Basalt.

The United States Army Corps of Engineers (USACE) encountered the Quincy

Diatomite deposit on the eastern side of Moses Lake/Potholes Reservoir while drilling

(Figure 1A). However, the diatomite in these drill cores is less than 1 meter thick and is thermally altered by the basalts, with nearly complete destruction of all diatom .

Overall, the Top Crude can be found up to 65 kilometers east of the Frenchman Hills, and up to 30 kilometers to the north and 15 kilometers to the south (Figure 1A).

Diatom Ecology

Diatoms are unicellular, golden , and are members of the division

Bacillariophyceae (Round et al., 1990). Their cell walls are composed of two valves, which occupy each side of the cell. These structures are made of opaline silica secreted by the diatoms, with one valve overlapping the other during the cell’s lifespan. Because

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diatoms cells, called frustules, are made of silica they are readily preserved in sediments, and are thus useful for paleoecologic reconstructions (Round et al. 1990).

Diatom species are differentiated as pennates or centrics, with the pennates being classified as araphid (without a raphe, which is a central slit-like perforation in the valve), monoraphid (raphe on one valve), and raphid (both valves have a raphe, motile) (Round et al., 1990, Williams, 1993). Pennate diatoms have rows of striae, which are composed of individual or double punctae. These striae and punctae, their number, and position and arrangement on the valve face are all used to differentiate genera and species.

The ecology and paleoecology of diatoms is based principally on species-specific preferences for certain environments and ecological associations. Diatom presence and abundance in a water body are in part related to varying preferences for particular limnologic conditions (Patrick 1977). Concentrations of metals and other non-nutrient elements may also act as limiting factors affecting diatom growth and reproduction.

Within a given lake, individual species of diatoms are observed in specific microhabitats within lakes, with these partitions being particularly well defined within the photic zone

(Bradbury and Forester, 2002). Paleolimnologic reconstructions rely on these observations, and use data sets of species assemblages accompanied by measured water to interpret changes in ecologic conditions through the sedimentary record of a given water body based on changes in floral assemblages.

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III. Methods

Sampling

Samples were taken from four localities, the Frenchman Hills, the Gorge

Amphitheater, Ancient Lakes Park, and Moses Lake (Figure 1a). Material from two of the localities (the Frenchman Hills and the Gorge Amphitheater) was obtained from drill cores provided by World Minerals from recently active mining sections. A third locality, the Quincy Ancient Lakes Park, is an outcrop in the Ancient Lakes Trail Campground, approximately 25 kilometers north of the Frenchman Hills. Material from the fourth locality is also from drill cores provided by the United States Army Core of Engineers, drilled on the north end of Moses Lake and Potholes Reservoir (Table 1). The Moses

Lake cores are extensively altered and did not produce diatoms, so only the first three localities are discussed in detail for this study. All samples and their corresponding cores/outcrops appear in Appendix 1.

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Table 1: UTM coordinates for all sample sites. Samples taken at each site are listed. All UTM coordinates are in NAD 83. Unit abbreviations are 4C, Four Crude, BC, Bottom Crude, and TC, Top Crude.

 +.*# -0# 32!0-.  +.*#'2#  +.*#,3+ #01 Q0412-11 Frenchman Hills 07AM05-17 to39 Q042-4 Frenchman Hills 07AM05-48 to 54 Q047-161 Frenchman Hills 07AM05-55 to 62 Pit Wall, section Frenchman Hills 07AM05-01 to 02 Pit Wall, section Frenchman Hills 07AM05-03 to 08 Pit Wall, section Frenchman Hills 07AM05-09 to12 Pit Wall, section Frenchman Hills 07AM05-13 Pit Wall, section Frenchman Hills 07AM05-14 to 15 Pit Wall, section Frenchman Hills 07AM05-16 Q01-20-9 Gorge Amphitheater 07AM05-40 to 45 Q01-20-15 Gorge Amphitheater 07AM05-46 Q03-16-23 Gorge Amphitheater 07AM05-63 to 64 Q01-9-1 Frenchman Hills 07AM05-65 Pit Wall, section Gorge Amphitheater 07AM05-47 Quincy Ancient Lakes Park ALP 07AM07-01 to 09 99BW15 Moses Lake 07AM08-06 to 08 00BW06 Moses Lake 07AM08-04 to 05 00BW14 Moses Lake 07AM08-01 to 03

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-,#   -++#,21 11 T 5201015 280230 Complete core, all 3 units present 11 T 5207445 273440 Complete core, only TC present, processed but not counted Complete core, all 3 units present. Only TC and 4C sampled, not 11 T 5208435 275550 counted 11 T 5206895 278885 Reconnaissance samples, processed but not counted 11 T 5207168 278520 Reconnaissance samples, processed but not counted 11 T 5207142 278511 Reconnaissance samples, processed but not counted 11 T 5207138 278539 Reconnaissance sample, processed but not counted 11 T 5207130 278516 Reconnaissance samples, processed but not counted 11 T 5207099 278511 Reconnaissance sample, processed but not counted 11 T 5214030 274461 Complete core, TC only present 11 T 5214230 274200 Sample of palagonite pillow breccia, not counted 11T 5215855 275525 Sample of palagonite pillow breccia, not counted 11T 5215930 275225 Sample of palagonite pillow breccia, not counted 11 T 5214427 274038 Reconnaissance sample, processed but not counted 11 T 5226465 277840 South facing outcrop along the northern trail 11 T 5228270 325600 USACE Core, thermally altered, not counted 11 T 5228840 322735 USACE Core, thermally altered, not counted 11 T 5228745 325960 USACE Core, thermally altered, not counted

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Lab Work

Samples were cleaned by placing approximately 1-3 grams of diatomite in 1-dram glass vials using 3% hydrogen peroxide for removal of residual clay particles. A diluted slurry derived from the washed sample vials was used to coat 0.1mm thick cover slips, which were then mounted onto 1.0mm thick, standard-width glass slides using either

Zrax© mounting medium (refractive index ~1.7) or Piccolyte (refractive index at least

1.52). At least two slides of each sample were made (yielding over 200 slides total).

Slides were examined using an Olympus BX-51 microscope at 400X and 1000X magnification; light micrographs were taken using an Olympus DX-71 microscope camera with Olympus software. Specimens were also examined using a liquid nitrogen- cooled Jeol JSM 840A Scanning Electron Microscope equipped with Fissions Kevex digital beam control interface and software for photography.

Frustule point counts were done in a linear transect on each slide for a core from the Frenchman Hills (Q0412-11) and for the samples from the Ancient Lakes Park.

Frustules were only counted if they were greater than 50% intact. Point count data are presented in Appendix 3.

Error for the relative abundance counts were determined by comparing the two slide counts per sample. Differences between the two counts ranged from 0.3% to 7% among all samples, with most samples having an error range between 0.5% and 2%

(Appendix 4). In examining the percent differences between individual species in each of the two counts, the largest variation was in counts of Aulacoseira granulata, the most abundant species throughout the diatomite.

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Data Analysis

Point count data were analyzed using the program R, version 2.9.1, available in the public domain. Extremely rare taxa (those with less than 2% total occurrence in any sample, and those with less than two occurrences over all samples counted) were removed from the data set in order to create model that reflected the dominant taxa present. NonMetric Multidimensional Scaling (nMDS) was used to analyze counts from the Frenchman Hills and the Ancient Lakes Park. NonMetric Multidimensional Scaling is an unconstrained ordination useful as an alternative to Correspondence Analysis.

Matrix calculations used in this technique are less sensitive to non-linear relationships and discontinuous distributions, making the method more likely to display the real differences between species and samples and provide more robust results for interpretation of data (Clarke and Ainsworth, 1993, McCune and Grace, 2002). Samples within the dataset were given the sample numbers used during collection, and species were identified using letters starting with D and continuing through to double letter denotation (Table 2). Data from nMDS calculations were plotted against factors 1 through 3 for analysis of which species and samples group together and how the species and samples vary between each other. Samples were also analyzed with a cluster analysis using Ward’s Minimum Variance method for determination of relatedness among the samples. The results of this cluster analysis were then compared to those from nMDS for interpretation of relatedness of samples and taxa.

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IV. Observations

Stratigraphic variation within units observed in Frenchman Hills

Species-level analysis confirms the three units that comprise the Quincy

Diatomite deposit have distinctive floral compositions with lithologic properties linked to variations in species composition. Analysis of mean valve width (Appendix 5) shows the upper intervals of the Top Crude have the smallest mean grain size, averaging approximately 7 μm. The Actinocyclus-rich lower interval of the Top Crude and the entire Bottom Crude, have the largest mean grain size, and the Bottom Crude also has the largest valves in the entire deposit (~ 50μm wide). Four Crude mean valve size is similar to the Top Crude. Variations in valve size can be directly correlated with the macroscopic lithologic properties of the diatomite, and these variations produce three visually distinct units with distinctly different feels to the touch. These major floral changes observed in the point count data correspond to the discernible visual and textural differences observed in the field.

The Frenchman Hills Flora

In the Frenchman Hills core, 75 species were counted (Appendix 3). The lowest unit, the Four Crude, contains 43 of these 75 species and is dominated by centric diatoms.

Aulacoseira granulata (Ehrenberg) Simonsen is the most abundant, ranging from 30.0-

79.0% of specimens counted in a given sample (Figure 4), with a mean of 48% for all samples in the Four Crude. Aulacoseira canadensis (Hustedt) Simonsen is also abundant in the lowest two samples (39.3% and 25.0% respectively), but its abundance decreases to less than 1% up section (Figure 4). Actinocyclus species abundance is highly variable,

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ranging as high as 55.3% within a single sample, and araphid pennates are the most abundant pennate form, with Staurosira construens var. venter (Ehrenberg) Williams and

Round being the most common. The araphid pennate to centric ratio (A:C ratio), useful for generalized paleobathymetric interpretation (with higher values of araphids corresponding to more shallow bathymetries), ranges from a low 4:294 (0.014) to 49:239

(0.21) in the Four Crude, with a median ratio of approximately 16:281 (0.057). This ratio is controlled largely by localized abundance fluxes in S. construens v. venter (Figure 4).

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Figure 4: Stratigraphic plot of relative abundance of the five most abundant species and the araphid pennate to centric diatom ratio at the Frenchman Hills, core 0412-11. Percent abundance is plotted for each species within each sample. Thickness is in meters below the top of the core.

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The Bottom Crude is dominated by Ellerbeckia baileyi, A. granulata, and S. construens v. venter (Figure 4), and overall species diversity is greatest in this unit (69 species counted from the unit) (Appendix 3). E. baileyi typifies this unit, with a range of

38-74.7% of species counted. A single sample near the base of the unit, where material is poorly preserved and possibly reworked (sample 07AM05-31, depth 14.7 ft), is composed primarily of E. baileyi fragments; specimens preserved well enough to be counted were thus only those that were exceptionally well preserved. In this sample, the numbers of S. construens v. venter were highest (23.7%). Overall, abundance of E. baileyi decreases up section to < 5% in the uppermost sample (Figure 4). Species of

Planothidium also occur nearly exclusively in this unit in moderately low abundances of

(2-4%) throughout the Bottom Crude (Appendix 3). The A:C ratio of the Bottom Crude ranges from 18:280 (0.064) to 139:124 (0.89), with a median ratio of 44:248 (0.18).

The Top Crude is dominated by A. granulata, with localized zones of increased abundances of Actinocyclus and A. canadensis. Species diversity in the Top Crude is low

(17 species counted over all samples from the unit) (Appendix 3); centric diatoms dominate throughout, ranging from 73.0 – 100.0% relative abundance (Figure 4). The genus Actinocyclus (in particular, Actinocyclus motilis Bradbury and Krebs) is most abundant in the lowest parts of the Top Crude. High Actinocyclus abundance also co- occurs with a slight increase in pennate abundance. A. canadensis species counts increase in the middle portions of the Top Crude, and the upper 2 meters are dominated by A. granulata (Figure 4). Combined, the three species of Aulacoseira observed account

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for 99.7% of all specimens counted in the upper 3 meters of the Top Crude. This succession of centric diatom dominance from A. motilis to A. canadensis followed A. granulata creates three florally distinct intervals within the unit. The rare pennates that were counted are most commonly araphid, and are usually species of

Tetracyclus, Synedra, or Meridion. Diatomite samples were also taken approximately 2 cm above and below the mafic air-fall tuff located near the base of the Top Crude (Figure

3) and no floral changes were observed across the air-fall tuff. A:C ratio is consistently low for the entire Top Crude; most often the ratio is 0:300 (100% centrics), with only the two bottom samples having any araphid pennates counted. The lowermost two samples have a ratio of 54:242 (0.22) and 15:272 (0.055) respectively (Figure 4).

The Ancient Lakes Park (ALP)

The Top Crude is the only unit exposed at the ALP, which is the northern most sample locality in the western area. The thickness of the Top Crude is unknown at this location because the base is covered but it is estimated to be no more than 6 m thick based on outcrop size and the projections of basalts exposed in nearby areas.

The Top Crude at the ALP is Aulacoseira-dominated, but it has significantly higher species diversity (39 species counted) compared to the more southerly locations.

It displays a heterogeneous floral assemblage with no apparent stratigraphic succession of species. Samples are dominated by A. granulata, A. canadensis, and Fragilariforma virescens (Ralphs) Williams and Round, with a nearly even mix of A. granulata and A. canadensis being most common; A. granulata is the most abundant centric diatom, with

A. canadensis being second most abundant at the ALP (Figure 5). Combined, species of

Aulacoseira account for 56.3-95.0% of all species counted.

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Araphid pennates are very common at this locality. F. virescens is the third most abundant species at the ALP, displaying peak abundance in the middle samples of the outcrop (Figure 5). Of the remaining araphid pennates commonly found, Meridion circulare (Greville) Agardh is next most abundant, and species of Tetracyclus, particularly Tetracyclus linearis (Ehrenberg) Grunow occur commonly at the base of the section. The ALP section also has higher abundances of raphid diatoms compared to the

Frenchman Hills. In particular, the genus Gomphonema accounts for up to 11.3% of all specimens counted (Figure 5). Species of Eunotia, Pinnularia, and Navicula are also found more commonly here than at the other, southerly sampling localities.

The A:C ratio of the Top Crude at the ALP differs significantly from the Top

Crude at the Frenchman Hills. The Top Crude in the Frenchman Hills is dominated by centric diatoms, but the Top Crude at the ALP has a reduced abundance of centrics, and the Actinocyclus –A. canadensis – A. granulata succession is not observed (Figure 5); the

A:C ratio ranges from 4:295 (0.014) to 74:171 (0.43), with a median ratio of 31:258

(0.12) (Figure 5). This range in A:C ratios at the ALP is actually most similar to that in the Frenchman Hills Four Crude unit.

Sample 07AM07-04 (depth 1.0m) is an anomalous sample, with many cells exhibiting internal valves and resting spores. M. circular shows these internal valves clearly, and several Aulacoseira and some unidentified centric species (Actinocyclus sp.?) exhibit resting stage cells and initial cells (Plate 19). This sample also shows an anomalously low abundance of A. canadensis, and many samples of what is interpreted to be A. granulata show diagenetic alteration similar to that seen by Thomas and Gould

(1981). These resting cells and internal

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valves occur exclusively in the ALP with their peak abundance in this interval, and they are not seen in the southerly sampling locations. Counts of these resting spores show they represent approximately 41% of the diatoms in the sample (Figure 5).

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Figure 5: Stratigraphic plot of relative abundance of the five most abundant taxa and araphid pennate to centric diatom ratio at the Ancient Lakes Park outcrop. Thickness is in meters above the base of the section.

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NonMetric Multidimensional Scaling Analysis Plots of the point count data through three data factors generated via nMDS produced a non-metric R2 fit to the data of 0.996, and a linear R2 fit of 0.984. The fitting of the nMDS shows statistical significance to the floral changes between the three units in the Quincy Diatomite deposit, confirming the distinct floral assemblages observed in the three stratigraphic units.

The three units, Four Crude, Bottom Crude, and Top Crude, are widely dispersed along factor 1, but they are much more closely distributed along factor 2 (Figure 6A, B).

The Four Crude (AM05-33 to AM05-39) and the Top Crude (AM05-19 to AM05-26) units group together tightly on the left side of the graph. The tight clustering of the Top

Crude and Four Crude samples in the nMDS plots do not resolve distinct floral intervals.

The Bottom Crude samples (AM05-27 to AM05-32) are the most out-lying, and are more distinctly separate from others in the upper portions of the graph. These samples show a strong association with species found dominantly in the Bottom Crude (Figure 6A), such as E. baileyi, S. construens v. venter, and species of Planothidium and Gomphonema.

Several species associations are noted for potential ecologic significance (Figure

6A). Species grouping near the samples of the Bottom Crude include E. baileyi. While

P. brevistriata is not a benthic species (planktonic, possibly littoral according to Abbot and Vanlandingham 1972), it and its varieties (P. brevistriata v. subcapitata) more strongly correlate with shallow water benthic species found commonly in the Bottom

Crude. Planktonic species, such as A. granulata, A. canadensis, and all species of

Actinocyclus group together near the Top Crude and Four Crude units.

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One sample in particular from the ALP, AM07-04, is clearly separated from the other groups along factor 2 and is interpreted to represent a distinct ecological association (Figure 6B). This sample contains the relatively high abundance of resting spores and altered Aulacoseira cells. It is also associated with increased abundances of

M. circular (often with internal valves), Eunotia curvata (Kützing) Lagerst, and species of Gomphonema.

When species and samples are plotted against factor 1 and factor 3 (Figure 6C, D) a wider dispersion within the data is visible. Species associations appear very similar to those observed in plots of factors 1 and 2, with E. curvata, E. baileyi, and species of

Gomphonema again being associated with the Bottom Crude, and the planktonic species showing strong association with Top Crude samples. ALP samples again cluster near samples from the Four Crude and Top Crude units. Broadly, the Bottom Crude samples are still distinctly separated from other samples, with the discordant sample from the ALP

(AM07-04) also grouping with the Bottom Crude.

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Figure 6: Nonmetric Multidimensional Scaling analysis (nMDS) plots of species and samples from the Frenchman Hills (core Q0412-11) and the ALP (samples underlined). Numbers of samples correspond to those shown in Table 2. 6A shows nMDS of species only, plotted with factors 1 and 2. Arrows represent sample vectors representing degree of similarity of samples. Samples and their associated species from the Bottom Crude are closed in the irregular box to show their clustering. 6B shows sample vectors with labeled arrows, plotted with factors 1 and 2. Longer arrows indicate samples accounting for more variability and more distinctive species compositions, while arrows near the origin of the graph show samples with many common species. Species positions seen in 6A are designated by dashes. Samples and their associated species from the Bottom Crude are enclosed in the irregular box to show their clustering. 6C shows species plotted against factors 1 and 3, and unlabeled arrows showing position of samples. 6D shows samples plotted against factors 1 and 3, with labeled arrows, and unlabeled species plotted by dashes.

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Table 2: Species occurrence list. Species letter designations are those used for correspondence analysis. Lower case “nc” indicates a species is observed but not counted, and thus not included in the nMDS.

Species Species CODE Actinocyclus motilis D Actinocyclus krasskei E Actinocyclus sp F Anomoeoneis sphaerophora BG Aulacoseira agassizi G Aulacoseira canadensis H Aulacoseira granulata I Aulacoseira granulata (?) diagenetic J Cavinula pseudoscutiformis BV Craticula sp. BH Cymbella cistula K Cymbella ventricosa L Diatoma anceps M Diatoma hyemale N Diatoma hyemale v mesodon O Diploneis ovalis BI Ellerbeckia balleyi P Eunotia curvata Q Eunotia pectinalis R Eunotia pseudopectinalis nc Eunotia pectinalis var minor BJ Eunotia veneris S

Pseudostaurosira brevistriata T Pseudostaurosira brevistriata var. elliptica BK Pseudostaurosira brevistriata var. subcapitata U Fragilaria capucina var. laceolata BL Staurosira construens V Staurosira construens var. venter W Staurosira construens var. binodis nc Fragilaria lapponica Y Fragilaria near lenoblei Z Fragilaria leptostauron AA Staurosirella martyi X Fragilaria pinnata var lancettula AB Fragilaria near socia AR Fragilaria vaucheriae BM

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Fragilariforma virescens AC Fragilariforma intortus AD Gomphonema acuminatum var montanum BN Gomphonema angustatum BO Gomphonema cholnokytes AE Gomphonema gracile BB Gomphosphenia grovei BR Gomphonema grunowii BC Gomphonema affine var. insigne AF Gomphonema dichotomum AG Gomphonema tenellum BQ Melosira undulata AH Meridion circulare AI Navicula acceptata AJ Navicula avenacea BS Navicula near insula BT Navicula pantoesekiama BU Navicula reimertes AM Navicula scutelloides BW Navicula seminuloides BX Neidium near iridis var. ampliatum nc Nitzschia amphibia AN Pinnularia near major BD Pinnularia nodosa BE Placoneis amphibola AK Placoneis anglica nc Placoneis near gastrum AL Placoneis elginensis nc Placoneis placentula nc Placoneis rostrata nc Achanthidium exiguum nc Planothidium conspicuum AO Planothidium ellipticum BY Planothidium lanceolatum BZ Planothidium rostratum AP Rhoicosphenia curvata CA Sellaphora near americana nc Sellaphora laevissima nc Sellophora bacillum AQ Stauroneis near acuta BF Stauroneis phoenicenteron nc Synedra tabulata var. fasiculata BP

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Tetracyclus ellipticus AS Tetracyclus lacustris AT Tetracyclus lata AU Tetracyclus lancea AV Tetracyclus linearis AW Tetracyclus polygibbum AX Tetracyclus williamsensii AY Reimeria sinuata nc

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Cluster Analysis

Cluster analysis of samples from the Frenchman Hills and the ALP produce clusters with sample associations similar to those observed in nMDS. Four distinct groups can be separated from the cluster analysis (Figure 7). One group is composed of a single sample; this sample from the ALP, AM07-04, is the anomalous sample observed in point counts and nMDS plots (Figure 6). It is most closely related to two samples from the lower Bottom Crude (AM05-30 and AM05-31), which also produce their own cluster.

Samples from the ALP are part of a cluster that includes the lowermost Four Crude and uppermost Top Crude samples. Samples showing the transition from the Bottom Crude to the Top Crude also cluster together in another group. These clusters show Four Crude and Top Crude samples associated in a similar manner to that observed in the nMDS plots.

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Figure 7: Cluster analysis of samples from the Frenchman Hills and the ALP used for point counts showing four primary clusters based on statistical similarity (distance) of species composition using the Wards linkage methodology. Samples from the ALP are underlined.

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V. Limnologic Interpretations

The ecology and paleoecology of the lacustrine environment are based principally on species-specific preferences for certain ecological conditions. Diatom presence and abundance in a water body are heavily influenced by varying preferences for particular limnologic conditions, including nutrient budget and other ecologic variables (Patrick

1977). Modern diatom ecology studies often use multivariate analytical techniques to correlate limnologic and ecologic variables to species assemblages within a water body and thus identify the ecologic preferences of individual species (Gasse, 1987, Starratt,

1987, Fitzpatrick et al., 2003, Telford et al., 2004). Many paleoecological studies on existing lakes that employ diatoms for paleolimnologic reconstructions, often going back

10 ka, have developed a modern calibration set to determine modern ecological associations. These studies then use quantitative analyses from the modern assemblage to describe how the paleoassemblages deviate, and the implications for how water quality parameters vary over time (Battarbee, 1986, Bennion et al., 1996, Bradbury and Forester,

2002, Lotter et al., 1997, Lotter et al., 1998). These studies demonstrate that diatoms have high sensitivity to water quality variables, and that the characteristics of and changes in dominant species and floral assemblages can be used to infer paleoecologic conditions in time and space.

Miocene paleoecology and paleolimnology is significantly more speculative than

Holocene reconstructions. Miocene lakes with no modern expressions present a unique challenge for paleolimnologic study, as back calculating water chemistry based on recognition of statistically significant ecologic gradients within the lake system is not

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possible for these deposits. Most Miocene species are extinct and do not have established optima or ranges of tolerance for ecologic variables, as do many extant species. Despite these limitations, inferences regarding Miocene paleoecology have been made in the published literature. In particular, Abbot and Vanlandingham (1972) published a compilation of previously published works on modern diatom ecology, and they attempted to apply this knowledge to a Miocene diatomite deposit. The previous works detailed the physiochemical preferences of several diatom species qualitatively.

These preferences included pH tolerances, dissolved salt tolerances (halobion spectra), preferences (limnophylic spectra), and nutrient level preferences (trophic spectra).

The published general assessments for species were compiled and assessed for agreement into a set of spectra as a way of interpreting the paleoecology of an extinct lake. The ecologic spectra of Abbot and Vanlandingham (1972) thus represent a way to qualitatively describe the ecology of the ancient Quincy Lake before the use of more recent statistical techniques for extrapolation of limnologic conditions.

Use of the spectra and dominant species as ecologic proxies is justified by the species assemblage observed at Quincy. While it is impossible to know how much the ecologic preferences of individual species has varied over time, many of the extant species in the Quincy diatomite deposit are morphologically identical to their living representatives. Further, the species assemblage itself is very similar to modern assemblages, with inter-species associations and relative abundances of dominant species similar to those encountered in existing lakes. Despite the lack of definitive knowledge regarding diatoms temporal consistency for specific ecologic parameters, the similarity of this diatom assemblage to modern lakes provides a basis for interpretation.

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Lake Nutrient Budget/Paleolimnologic conditions

Paleolimnological interpretations of the ancient Quincy Lake must rely heavily on information known about extant species within the Quincy diatomite deposit (48 species are confirmed to be extant), and on habitat associations inferred for genus- and species- level groups. In particular, these interpretations rely heavily on the extant species

Aulacoseira granulata, the most dominant diatom in the deposit, and they are supplemented with ecological data on several other species that have significant spikes in dominance over various stratigraphic intervals, including Staurosira construens var. venter. A. granulata averages 60.3% abundance in the Four and Top Crude, and it reaches a maximum abundance of 99% in the upper intervals of the Top Crude (Figure 4,

5). It is of slightly lesser abundance in the Bottom Crude, ranging from 13% to 39% relative abundance, with a mean abundance of approximately 26% (Figure 4). A. granulata is thus viewed as a primary proxy for the ancient Quincy Lake history, particularly for the Four Crude and Top Crude. It is also viewed as an important ecologic indicator for the Bottom Crude.

The Abbot and Vanlandingham (1972) spectra for A. granulata indicate the ancient Quincy Lake was alkaline, with a pH above 7, and that the lake was eutrophic.

These spectra also suggest the lake was oligohalobous, with a dissolved salt content less than 5‰. Spectra reported for S. construens v. venter and P. brevistriata also support an oligohalobous, alkaline lake (Abbot and Vanlandingham 1972) as does the presence of

Actinocyclus motilis (Bradbury and Krebs 1995). The spectra further suggest a high abundance (> 50%) of euplanktonic species (such as Aulacoseira or Actinocyclus), indicating the ancient Quincy Lake must have been greater than 15 meters deep during

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both Top Crude and Four Crude deposition. The autecology (the obligate ecologic environment of a particular species for subsistence) for A. granulata is further evidence of a productive lake with a sizable standing crop of phytoplankton; A. granulata’s autecology is that of a euplanktonic (obligate planktonic) diatom (Davey 1986, Sicko-

Goad et al. 1986). As such, A. granulata is an indicator of open water during deposition of sections of the stratigraphy where it is dominant. The near 100% abundance of euplanktonic species counts in the Top Crude suggests that this upper unit represents the deepest water stage for the ancient Quincy Lake. These qualitative assessments are further bolstered by more recent, quantitative studies based on transfer functions and measured limnologic variables of several lake parameters such as total phosphorous, hydrogen ion concentration, and (Bennion et al., 1996, Slate and Stevenson,

2000, Schelske et al., 2006).

Specific limnologic conditions for the ancient Quincy Lake can be approximated using several species as proxies, extrapolated from work with transfer functions on modern data sets. Total phosphorus (TP) is one important chemical variable in terms of lake ; it has been shown to affect phytoplankton growth, abundance, and floral assemblages, in addition to being the overriding factor affecting trophic status in a lake (OECD 1982). Bennion et al. (1996) provided a model for a TP transfer function for use in paleolimnology based on several data sets available via the European Diatom

Database. This transfer function, based on combined European datasets, accurately predicted TP levels when A. granulata and several fragilariform species (i.e. species from genera related and morphologically similar to the genus Fragilaria) (including S. construens v. venter) combine for over 40% of species counted. According to the

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Bennion et al. (1996) data set, the species assemblage, and the calculated projection, abundances like these would imply TP levels of approximately 140 μg/l, with increases in planktonic centric species abundance indicating increased TP levels. Bennion (1996) noted these levels correspond to eutrophic systems, with many European lakes that were measured having TP levels exceeding 200 μg/l. These values would equate to hypereutrophic conditions, as defined by the OECD (1982), where TP mean annual values 100 μg/l are considered hypereutrophic according to this classification scheme.

Based on the dominance of A. granulata and the extrapolated TP levels, the ancient

Quincy Lake is interpreted as having had an annual average TP level of at least 100 μg/l, and likely in the 140 μg/l range, placing the lake well in the range of a eutrophic to hypereutrophic system.

Total hydrogen ion concentration (pH) is another important ecologic variable.

Gasse (1986, 1987) published a transfer function for calculating pH values based on an equation accounting for the percent abundance of a species, the mean pH parameter for that species, and a regression coefficient specific to each species. Published mean pH preference values for A. granulata include 7.5 by Gasse (1987), 7.7 by Bennion et al.

(1996) and 8.2 by Lotter et al. (1998). The dominance of A. granulata therefore indicates a slightly alkaline lake throughout the lake history. Averaging the published values for A. granulata yields a mean pH parameter of 7.8; using this value in the equation from Gasse

(1986, p.159) then leads to the calculated average pH estimation of 7.7 for the ancient

Quincy Lake. Variability from this interpreted slightly alkaline value in the Four Crude and Bottom Crude is hypothesized to have been minimal, because of the dominance of A. granulata throughout these units.

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S. construens v. venter may be a also useful proxy for pH in the Bottom

Crude. Jones and Birks (2004) published a pH reconstruction based on calibration from several modern European lakes using a Weighted Averaging-Partial Least Squares (WA-

PLS) regression model that incorporated S. construens v. venter. They found that the reconstructed pH of the lake was above 7 (between 7.2 and 7.6) when S. construens v. venter was at its highest abundances. Jones and Birks (2004) also published the pH optima of all species found in their deposit, and the species tolerance of pH fluctuations based on their regression calculations; S. construens v. venter has pH optima of 7.6 with a tolerance of ±0.4 off that optima. Hydrogen ion fluctuations during Bottom Crude deposition are therefore interpreted to have been minimal, and the interpreted pH of the

Four Crude and Top Crude appears consistent with that of Bottom Crude deposition as well.

Dissolved silica concentration is also considered an important nutrient for diatom growth and development. A. granulata is most commonly found in waters with high silica content, and Kilham (1971) reported the average dissolved SiO2 of lakes is13.4 mg/l (ranging from 1.5 to 28mg/l in the published literature) when this species occurs as the dominant diatom in the water column. Optimal growth for this species is reported at

5 – 10 mg/l (Kilham and Kilham 1975), and is interpreted to be a proxy for eutrophic conditions (Kilham 1971). Modern lakes are highly variable in dissolved SiO2 content, ranging from 0.1 to over 60 mg/l, with a global average of approximately 12 mg/l (Horne and Goldman 1994). Based on the minimum values indicated by Kilham and Kilham

(1975) and Kilham (1971), dissolved silica levels in the ancient Quincy Lake were likely in the 5mg/l range, high enough to permit abundant A. granulata development in the

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deeper areas of the lake around the Frenchman Hills, and to permit sustained dominance of this species in the water column.

Gasse (1987) used a pH transfer function and estimations of paleosalinity to create several fossil assemblages associated with generalized lake conditions in modern

African rift lakes. Gasse found that assemblages dominated by A. granulata, with co- occurring Aulacoseira agassizi (Ostenfeld) Simonsen, Staurosira construens (Ehrenberg)

Williams and Round, and Pseudostaurosira brevistriata (Grunow) Williams and Round, typify the fossil assemblage that Gasse called 1D, indicative of a wide, tropical, shallow lake. Several other rare raphids known to occur in the deposit are also marker species for this assemblage classification, including Placoneis gastrum (Ehrenberg) Mereschkowsky,

Navicula seminuloides Hustedt, and Navicula scutelloides Smith. The African lakes that fit this classification (and thus the studied lakes diagnosed as having similar chemistry from their diatom flora) would have had a dissolved salt content < 0.5‰ and would be expected to be highly mixed and have high productivity. The interpretation of high productivity also fits with the sheer volume of diatomite (approximately 12 meters thick at some points), making this African rift assemblage a good analogue for the ancient

Quincy Lake.

In summary, the dominance of A. granulata through most stratigraphic intervals

(particularly the Four Crude and Top Crude), suggests relatively stable limnologic conditions with high TP levels and sufficient dissolved Si levels capable of sustaining a large euplanktonic population in the open water. The water would have been slightly alkaline, and the high nutrient levels and abundant planktonic diatom population would have been part of a eutrophic to hypereutrophic lake, with very high productivity.

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Genus Level Variation within the Quincy Diatomite deposit and its Ecologic

Significance

The alternating dominance of Actinocyclus and Aulacoseira in the Top Crude may represent variation between monomictic (one mixing event annually) and polymictic

(perennially mixed) lake circulation. Bradbury and Krebs (1995) interpreted alternation in the dominance of these two genera to be a function of circulation patterns in a monomictic lake, where Actinocyclus would dominate before mixing when the lake was stratified, and Aulacoseira would dominate during mixing. Aulacoseira would also dominate in a polymictic lake because of the continuous mixing. Using these criteria, the ancient Quincy Lake is interpreted to have been monomictic during Four Crude deposition, indicated by greater abundances of Actinocyclus (Figure 4). The Bottom

Crude was likely polymictic due to the dominance of benthic species like Ellerbeckia baileyi and the persistence of A. granulata throughout the unit. Based on the dominance of A. granulata in the intervals of peak Actinocyclus abundance, the ancient Quincy Lake would have become monomictic again during its expansion across the Quincy Basin, signaled by early Top Crude deposition. The lake would have then shifted to polymictic in the final stages of the Top Crude, when Actinocyclus became progressively become more rare (Figure 4), and Aulacoseira continued to dominate. These Top Crude mixing stages are interpreted to have minimally changed the water chemistry (as indicated by the continual dominance of A. granulata), and are thought to have shifted rapidly between depositional units as no gradational changes are observed in either the lithology or the flora.

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Paleoecology during Bottom Crude deposition

The Bottom Crude plots separately on the nMDS, it indicates a separate paleoenvironment (Figure 6). There is reduced abundance of A. granulata is low (10-

39%) in the Bottom Crude, and E. baileyi is dominant, averaging 34% abundance, and locally ranging as high as 75%. As with the other species in this genus, E. baileyi is a considered a benthic centric diatom (Crawford and Sims, 2007). This interpretation appears substantiated by the grouping of E. baileyi with other benthic diatoms in the nMDS plots (Figure 6b, d), such as Staurosira construens v. venter, which has an abundance increase and ranges 8 – 24% in the Bottom Crude. Colonization of benthic sediments newly exposed in the photic zone has been observed by Bennion (1995), where colonization of the bottom sediment created dense diatom growths in artificial water bodies. Further, fragilariform species (in particular, S. construens v. venter) were observed lying directly on the surface sediment by Bennion (1995), and it is hypothesized

E. bailey would have likely done the same

The A:C ratio in the Bottom Crude is also higher to an average of approximately

17% araphid pennates. Stockner (1967, 1971) demonstrated that, in general, as eutrophy occurred the proportion of araphid diatoms increased relative to centrics. Stockner further showed increasing numbers of S. construens directly correlate to increasing lake eutrophy and higher productivity. Kilham et al. (1996) also reported that S. construens occurs in water with high Si:P ratios, and may be an indicator of eutrophic environments.

Based the higher levels of S. construens v. venter in this unit – which is the highest percentage of araphid diatoms in the Quincy Diatomite deposit – the published ecologic information for S. construens v. venter, and the knowledge that E. baileyi is a benthic

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centric diatom, the Bottom Crude is interpreted to indicate a period of shallower water with highest productivity.

Alternative interpretations of Bottom Crude are also examined. Subtle variations in salinity levels are known to affect diatom floral assemblages (Fritz et al., 1993). At the

Quincy deposit, salinity level fluctuations were likely minimal because of the equally minimal observed changes in the floral assemblage through time. Based on the optima of the known species at Quincy, large variations in alkalinity and pH are also unlikely. A change in some limiting nutrient or rare element in the water body was the more likely stimulus for E. baileyi to become dominant. Changes in the nitrogen to phosphorus (N:P) ratio or the Si:P ratio have been shown to drive floral shifts (Brugam, 1983, Schelsky et al., 2006, Stoermer, 1986). However, in the absence of a modern calibration, this idea cannot be tested in the Quincy Diatomite deposit.

ALP proximity to littoral environment and fluvial influences

The Ancient Lakes Park contains a larger littoral species component than the Top

Crude at the more southerly localities, and this may indicate either the ALP is reflective of the distal littoral environment. Planktonic centric diatoms still represent a mean of about 87% relative abundance at this locality; however the ALP Top Crude contains larger pennates such as Fragilariforma virescens, and a marked increase in epiphytic taxa, such as Gomphonema spp. (Figure 5). This contrasts with the Top Crude in the

Frenchman Hills, which is nearly devoid of pennates. Evidence for this location having been ecologically transitional comes from the geographic location and different floral assemblages (biofacies) shifts observed in the ALP. From a geographic standpoint, the

ALP is the most northern possible outcrop in the western side of the modern Quincy

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Basin and is hypothesized to be close to the littoral zone at the northern extent of the basin. While the Quincy Diatomite deposit does have one outcrop in a more northerly location more than 60 km to the east, this outcrop is only 6 km farther north.

The position of the Columbia River as a potential fluvial influence to the ALP may have provided significant ecologic impacts. Evidence has been presented to suggest the Columbia River has occupied the same approximate position since the middle

Miocene as it does today (Reidel et al., 1987, Martin, 1989, Martin, 1991). If this is the case, the Ancestral Columbia River would be projected to be the western boarder of the deposit, as is observed in outcrops (Figure 8). Flow from the river is not thought to have directly fed the ancient Quincy Lake due to the marked lack of clastic sediments in the diatomite. However, tributary inflow from surrounding streams near the ALP could have had a significant ecologic impact, including more variable local water chemistry, differential flow rates, and providing a source for periphyton to be washed into the ancient Quincy Lake. Lithologically, the ALP does have a slight increase in the amount of clay, which is visible during initial sample processing. It is also slightly more indurated than other Top Crude samples from the southern sampling locations. The slightly higher clay component at the ALP may be attributed to suspended load influx from a tributary system surrounding the Ancestral Columbia River and feeding the ancient Quincy Lake. Ultimately, without observation of a fan complex inter-fingering with the diatomite deposit, the true source of water and sediment influx is not fully understood and is perhaps slightly beyond the scope of this work.

The abundance of Aulacoseira canadensis is increased at the ALP and may indicate a distal littoral zone with more turbid conditions. Kilham et al. (1986) discussed

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pore sizes in the genus Aulacoseira and the corresponding light requirements. Their findings suggest an inverse relationship between the two variables. Littoral environments are more likely to be shaded by surrounding vegetation and can be highly turbid as wave motion may continuously disturb the sediments; phytoplankton would therefore receive less light than open water forms and are more likely to be disturbed if settling out of the water column occurs. A. canadensis has the largest pores of the three Aulacoseira species in the diatomite and is also heavily silicified. As such, it may have preferred environments where water circulation and increased sediment disturbance would have better kept the species in suspension compared to more open water.

It is worth noting that A. canadensis is not used here as a direct proxy for water chemistry because more information regarding the specific preferences of the species is needed. By inference and the strong association with A. granulata, A. canadensis is interpreted to be a slightly alkaliphilous species tolerant of eutrophic conditions. The previously noted relative abundance increase of this species in the Top Crude does not seem explainable via low pH levels, particularly since A. granulata still comprises more than 50% of species observed in most samples, despite the increased A. canadensis abundance (Figure 4). This is in contrast to the findings of Bahls et al. (2009). It is suggested the morphologic similarities observed by Bahls et al. (2009) do not necessarily equate to similar pH optima, and the unknown chemical preferences of the species prevent its use as a direct proxy for chemical parameters.

The ALP biofacies shows local indications of a stressed habitat. Several specimens observed have what are interpreted to be resting stages and internal valves

(Plate 19), particularly in the 1.0m interval, sample AM07-04 (Figure 5, Appendix 3).

52

These heavily silicified specimens may be indicating variable ecologic conditions, with possible discontinuous water supply, prompting the diatoms to enter resting stages.

Lacustrine waters in a shallower environment or those influenced by surface flow inputs are inherently more turbulent, and diatom frustules settling to the bottom or being washed in from a fluvial system may respond as if stressed. Regardless of whether some shift in local water level occurred, or some change in local water chemistry was induced (perhaps by tributary flows feeding the lake), it would have stressed the local diatom population.

The highest abundance of attached diatoms occurs in this interval (Figure 5), so some change in the local environment must have occurred to permit increased abundance of these benthic diatoms, while stressing the planktonic specimens enough to produce a depositional interval with relatively higher resting cell abundance. It is also important to note that species found in this particular interval, and in sample (AM07-04) itself, cluster with samples and species from the Bottom Crude in nMDS plots of factors 1 and 3

(Figure 6C, D), indicating a close association of these species and samples along an ecologic gradient. Observed increased abundance of resting cells and internal valves is thus interpreted to be from some physiochemical stress in local lake environment, producing this isolated abundance of resting forms in an otherwise homogenous microenvironment.

Basin Development /Lake Stage Development Model

Based on these interpretations of the depositional stages of the Quincy Diatomite deposit, it is possible to develop a model of basin development for the ancient Quincy

Lake. This model is based on the model presented by Houseman (2006), the modern

53

geographic position of the diatomite, the Columbia River Basalt flow geology, and tectonic models of the area presented in the literature.

The thickness of the Quincy Diatomite deposit in the Frenchman Hills, particularly the presence of two units not seen in any other location (the Four Crude and

Bottom Crude), indicates the western section of the Frenchman Hills anticlinal ridge was near the basin depocenter for the ancient Quincy Lake. The Four Crude and Bottom

Crude units commonly, but not always occur together, and these two units are interpreted to represent the early footprint of the ancient Quincy Lake. The northwesterly orientation of sediments in the Frenchman Hills, visible in 3-D modeling of the subsurface, indicates the source of the lower two diatomite units were in close enough proximity to form a small, northwest-oriented, interconnected pocket lake system with an irregular footprint in the south western portions of the Quincy Basin. This small system would have persisted during Four Crude and Bottom Crude times, and, based on modern deposit geography from drill cores, it would have covered an area approximately 25 km2 in total size (Figure 8A).

Locations of Top Crude outcrops indicate a broad lake that would have covered an area over 2600 km2, shallowing to the east, and deepening to the west. The thickening of the Quincy Diatomite deposit to the west is also consistent with a northwest-trending

Wanapum Basalt flows discussed above (Schminke, 1967, Hooper, 1997). It is therefore plausible that tectonically driven basin subsidence from the progressive development of the YFB was a contributing force driving the ancient Quincy Lake’s expansion and the subsequent deposition of the Top Crude across an area over 2600 km2 at it peak distribution (Figure 8B).

54

The location of the Ancestral Columbia River is also indicative of basin geometry. The Columbia River is considered to be at the western limit of the diatomite because the deposit is not seen in exposures west of the river. The Ancestral Columbia

River drainage appears to funnel the Roza basalts towards a more westerly direction, allowing for further propagation towards the west with each successive flow (Martin,

1991). Further, evidence can be found in basalts pooling near the southern section of the

Quincy Basin at the Sentinel Gap of the Saddle Mountains (Reidel, 1984, Martin, 1989), indicating flow of the river would have continued along its present southerly course. If this flow geometry is correct, the thickening of Quincy deposit corresponds to a gradational basin deepening towards the Columbia River, with the river being the western limit of the diatomite.

55

56

Figure 8: Modeled extent of the ancient Quincy Lake (AQL) during developmental stages. Figure 8A shows the expected expanse of the lake during deposition of Four Crude and Bottom Crude based on deposit data from drill core. Lake outlines hug the margins of the encountered Four Crude and Bottom Crude occurrences. Figure 8B shows the projected outline of the AQL during Top Crude deposition. Lake outline is based on known outcrop localities or drill core data, and the lake is modeled to have not filled the entire Miocene Quincy Basin. Red stars show sample locations for this study.

57

The timing of episodic uplift of the Frenchman Hills anticline during this time frame can also be constrained by the sedimentologic features of the units within the

Quincy Diatomite deposit. The Four Crude and Bottom Crude appear massive in outcrop, with occasional laminations being visible in areas where chert lens formation is particularly pronounced and the laminations have become silicified. There is no gradational shift in the point count data between the Four Crude and Bottom Crude, with the transition instead being dramatic and abrupt. There is also a chert horizon in between the Four Crude and Bottom Crude units in nearly all localities. Based on the abrupt floral turnover, a clear separation of the units stratigraphically, and the occurrence of the Four

Crude and Bottom Crude together with apparent minimal variation in lake footprint based on drill core data for these two units, it is hypothesized a pulse of local uplift of the

Frenchman Hills (Reidel 1984, Martin, 1991, Hooper, 1997) occurred before overall lake expansion. This small, localized uplift combined with high productivity sedimentation, would have raised to the bottom of the ancient Quincy Lake enough to create a photic zone inclusive of the lake bottom. This would have resulted in the dominance of the benthic E. bailey in the Bottom Crude, where it would have been able to colonize bottom sediments in these more shallow parts of the lake.

The dominance of centric diatoms through most of the Top Crude, suggests rapid lake expansion, which would have permitted more open-water forms to dominate the flora. This expansion would have occurred due to a more regional subsidence (Martin,

1991, Hooper, 1997) of the Quincy Basin, with variability in bathymetry providing a potential explanation for the anomalously high levels of pennates at the ALP. Water to

58

fill the basin is hypothesized to have come from the surrounding watershed, though specific inlets are not identifiable, as sedimentologic indicators of fluvial inlets (i.e. deltaic deposits) have not been observed in outcrop or drill core. Regardless of the source, the dominance and widespread distribution of the Top Crude indicate a basin- wide expansion of the lacustrine environment.

Though the diatom flora in the Top Crude unit suggests lake expansion and apparent deepening, there is also sedimentologic evidence for continued localized growth and uplift of the Frenchman Hills Anticline in the form of gravity flows and paleosols.

Gravity flow deposits may represent an over-steepening of the slope by continued localized growth of the Frenchman Hills anticline during Top Crude deposition. The paleosol developed at the top of the Quincy Diatomite deposit in Frenchman may also be indicative of localized shallowing of the lake and exposure of sediments due to growth of the anticline. Diatomite is observed as being incorporated into the paleosol, and the paleosol was baked during emplacement of the overlying Priest Rapids Basalt flow.

These relations indicate that soil formation (and hence local uplift of the anticline during overall basin subsidence) took place prior to complete burial of the Quincy Diatomite deposit by Wanapum Basalts.

VI. Summary and Conclusions

The Quincy Diatomite deposit is dominated by planktonic diatoms through most of its stratigraphy, with geographic and stratigraphic variability documented among the most abundant species. Stratigraphic variations in flora help define three units within the diatomite; from oldest to youngest, they include the Four Crude, Bottom Crude and Top

59

Crude units. Aulacoseira granulata is the most abundant taxa throughout, and serves as the principal proxy to characterize the lake system. A. granulata is supplemented by information available for Staurosira construens var. venter, the most abundant pennate. Using these species as ecologic proxies, an overall model of development of the Quincy Basin and the ancient Quincy Lake can be summarized.

Ancient Quincy Lake initially formed on relatively uneroded basalt of the Roza

IIa flow. This initial stage of the lake, represented by the Four Crude, was restricted to areas in the southern part of the basin, including the western part of the Frenchman Hills.

The lake formed a series of small, pocket lakes along a northwest trend on the surface of the basalt. These pocket lakes were monomictic, eutrophic, slightly alkaline, and, based on the dominance of A. granulata, likely deeper than 15 meters.

A shallowing of the pocket lake system, induced via tectonic activity or reduced inflow, marks middle lake stage history, is represented by the Bottom Crude and typified by the benthic taxon E. baileyi. These newly more shallow lakes are interpreted to have had depths of 10 meters or less, to have been polymictic due to their reduced bathymetry, but still eutrophic and slightly alkaline based on the marked increase of S. construens v. venter and persistence of A. granulata. By association, E. baileyi is interpreted as being an oligohalobous, high productivity, eutrophic diatom with high dissolved nutrient content requirements and alakaliphilic preferences (pH above 7), similar to A. granulata and S. construens v. venter.

Lake expansion away from the Frenchman Hills, represented by the Top Crude, would have been driven by inflow from surrounding watershed. This larger lake stage represents, the deepest interval during deposition of the Quincy Diatomite deposit

60

(greater than 15 meters), and would have maintained high productivity. At the ALP, at least one sample with high levels of resting stages (07AM-04) indicates localized perturbations in the environment, which created environmental stress on the flora. Late lake stage also shows variations in A. motilis dominance and A. granulata dominance, which may be explained by variation between a monomictic and polymictic system.

Eventually, the Roza IIb Basalt flowed through the northern areas near the Gorge

Amphitheater, signaling the effective extinguishing of northern parts of the ancient

Quincy Lake. Eventually the sediments of the Quincy Diatomite deposit were uplifted in the Frenchman Hills anticline, paleosol development occurred, and the Priest Rapids

Basalt capped the exposed sediments.

This study of the Quincy Diatomite deposit shows that it is possible to produce a plausible paleolimnologic model for Miocene diatomites by using extant species as proxies for paleoecologic information. Any model such as this benefits from related geologic studies providing constraints on local to basin-wide processes influencing the diatom flora. The integration of both floral and geologic data is essential for arriving at a comprehensive understanding of the setting, environmental and ecological conditions in which a diatomite deposit formed. These models also serve as a way to characterize specific areas and microhabitats within a deposit, and correlate locations for a more broad-scale understanding of the paleoenvironment represented by the diatomite. While some uncertainty is inherent to these interpretations due to the age of the deposit

(Miocene), the dominance of modern species within the assemblage lends credence to the proxies used and interpretations made. Further, the statistical methodologies employed

61

demonstrate the use of modern ecologic information and species proxies to interpret this fossil deposit yields information that is both plausible and statistically robust.

Though these data cannot be expected to fully represent the paleolimnology of the entire

Quincy deposit, this study shows that diatomaceous sediments, and in particular Miocene diatomites, are prime candidates for future paleoecologic investigations.

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Chapter 2: Systematic Paleontology

I. Systematics

These systematics use Patrick and Reimer v.1 and v.2 (1966, 1975) and Round,

Crawford, and Mann (1990) for terminology and identification of most species in the sampled material. Additional sources used for identification and specific terminologies are Bradbury and Krebs (1995) for the genus Actinocyclus, Williams (1996) for identification and terminology specific to Tetracyclus, and Crawford (2007) for identification and terminology specific to the genus Ellerbeckia. References used strictly for identification purposes include Schauderna (1983), Thomas and Gould (1981), and

Williams and Round (1987). All identification resources were used equally as principle literature sources. Permanent mount slides, remaining slurry material, and a portion of the dry, unprocessed samples will be stored at the University of Nevada, Reno,

Department of Geological Sciences and Engineering. A total of 84 species were identified within the deposit and are described below. The distribution for these species is not characterized beyond their occurrence in the Quincy Diatomite.

Division BACILLARIOPHYTA

Class:

Subclass: Coscinodiscophycidae

Order: Aulacoseirales Crawford 1990

Family: Aulacoseiraceae Crawford 1990

Genus: Aulacoseira Thwaites 1848

Aulacoseira agassizii (Ostenfeld) Simonsen 1979 63

Plate 1 number 1-9

Basionym: Melosira agassizii Ostenfeld 1908

Synonyms: Melosira schroderi Woloszynska 1914

Description: Valve height is always less than valve diameter, with height ranging 5 –

10μm, and diameter ranging 12 – 18μm. Striae are straight and continuous across

epi and hypo-valves and linked valves. Separation spines are robust and fairly

uniform in length. No single separation spine extends across the entire linked

valve. Punctae are polygonal. Ringleiste is strongly developed and is observable

as a thickly silicified line when focusing through the valve in girdle view.

Auxospore-type cells terminating cell chains are common for this species and are

often the most diagnostic.

Type Locality: Victoria Nyanza, Central Africa.

Remarks: Similar in appearance to A. granulata, however the separation spines are not as

long, and are more numerous. This species is also most commonly observed with

its auxospore in a chain of three to five valves. Vanlandingham (1964) noted

some A. agassizi frustules appear very similar to A. granulata, and in fact named

some of them A. granulata near agassizii. He also speculated this might represent

a continuum of variation with A. granulata on one end and A. agassizi on the

other. Specimens assigned to A. agassizii are restricted to those with greater

width to diameter ratios, and short, robust separation spines.

Distribution: Seen most commonly in the Frenchman Hills, usually in the Top Crude. It

becomes more rare moving north with sampling localities, being rare in near the

Gorge Amphitheater and nearly absent in the Ancient Lakes Park. 64

Aulacoseira canadensis (Hustedt) Simonsen 1979

Plate 2 number 1-10, Plate 19 number 6

Basionym: Melosira canadensis Hustedt 1952

Synonyms: none

Description: Valve diameter to height ratio ranges from 1:1 to 3:1, with rare exceptions

less than 1:1. Diameter has a range of 6 – 16μm and height ranges 9 – 23μm.

Separation spines are very small (~1μm) and of uniform length. Punctae are

commonly wrapped in a spiral pattern around the frustule, with this pattern being

more easily recognized in specimens with lower pore densities. Punctae density

is irregular, with spaces between even the most densely porous specimens.

Punctae are circular to ovate and are large for this genus (range 1-2m).

Ringleiste is strongly developed and easily visible when focusing through the cell

wall. Valves joining a complete frustule are hyaline at the junction, creating a no-

poroid area between the epi and hypovalves. This hyaline area is consistently 2 –

3μm in height.

Type Locality: Quesnel, B.C., Canada. Fossil. Harper, E. Oregon, USA.

Remarks: All specimens are heavily silicified. Increased abundance in the ALP, which is

interpreted to represent an area of more shallow water, may provide some

ecological information for this species. Further studies in the region could

provide more definitive results.

Distribution: This species occurs in all sampling localities, with greatest

abundance at the ALP, where it is may be most abundant diatom in certain

samples. It is also abundant in the mid-lower portions of the Top Crude, where it 65

has an abundance peak stratigraphically above the abundance peak of

Actinocyclus species.

Aulacoseira granulata (Ehrenberg) Simonsen 1979

Plate 1 number 10-12

Basionym: Gallionella granulata Ehrenberg 1843

Synonyms: Gallionella decussata Ehrenberg 1843, Melosira granulata Ehrenberg 1853,

Orthosira punctata W. Smith 1856, Melosira marchia (Ehrenberg) Ralphs in

Pritchard 1861, Melosira lineolata Grunow in Van Heurck 1882, Melosira

polymorpha subsp. granulata Bethge 1925.

Description: Diameter and length of specimens is highly variable with specimens of all

sizes co-occurring in any given sample. Valve height to diameter ratio is often

1:1 or greater, ranging up to 3:1, and with rare occurrences of 1:2. Height ranges

7 – 23μm and diameter 5 – 14 μm. Striae may be straight or show off-set sigmoid

pattern in girdle view and also exhibit robust separation spines which commonly

extend across the entire length of the opposite valve. Polygonal punctae are

visible through all specimens. Areola size varies from 0.5 – 1.5m in diameter

and the characteristic polygonal shape is always present. Ringleiste is very

prominent, and may become slightly undulate in specimens showing signs of

. Whole frustules show a non-poroid, hyaline section between epi and

hypovalves. This area is highly variable in height (1 – 4μm).

Type Locality: “Brasilen? Neufundland” (Ehrenberg 1843)

66

Remarks: There is large variation in the degree of frustule silicification, with heavily

silicified diatoms co-occurring with very lightly silicified specimens through all

sampling locations. Similar observations have been made by Stoermer et al.

(1981). In samples that exhibit signs of being reworked, with most specimens

broken, A. granulata shows signs of digenetic effects, including shrinking of the

pores similar to the observations by Thomas (1981). This is the only species

counted in every sample at every locality.

Distribution: Common across all sample localities and stratigraphic positions, this is the

most common and abundant diatom in the Quincy Diatomite.

Order: Melosirales Crawford 1990

Family: Melosiraceae Kützing 1844

Genus: Melosira Agardh 1824

Melosira undulata (Ehrenberg) Kützing 1844

Plate 4 number 1-3, Plate 20 number 6-7

Basionym: Gallionella undulata Ehrenberg 1840

Synonyms: none

Description: Specimens range 16 – 37μm in height, and 22- 35 μm in diameter,

with height to diameter ratio being nearly 1:1. Valves are finely striate on the

exterior, with a slight sigmoidal pattern, and are off-set across epi and

hypovalves, with the pattern becoming linear and straight across separation

valves. Areolae are of variable size and shape (subrectangular to circular), and

create canals through the heavily silicified valve walls, creating a segmented 67

appearance when focusing through the specimen. No separation spines are

present. Ringleiste appears as an “undulatory” wave pattern while focusing

through the specimen.

Type Locality: Uncertain

Remarks: Point counts often did not encounter this specimen, but at 400x magnification

several frustules are visible. The largest specimens come from the Frenchman

Hills, with the ALP specimens being smaller. All valves are heavily silicified and

most have picked up diagenetic staining not removed with the cleaning

procedures used. Single valves were most commonly encountered, with intact

chains being very rare.

Distribution: Rare across all samples, but found most commonly in the ALP.

Order: Coscinodiscales Round and Crawford 1990

Family: Hemidiscaeceae Hendey 1937 emend Simonsen 1975

Genus: Actinocyclus Ehrenberg 1837

Actinocyclus krasskei (Krasske) Bradbury and Krebs 1995

Plate 18 number 1-3

Basionym: Coscinodiscus miocaenicus Krasske 1934

Synonyms: Coscinodiscus miocenicus Krasske sensu Bradbury and Krebs 1995

Description: Valves circular and mostly flat, with minor concentric undulation visible in

the largest specimens. Diameter ranges 15 – 45m. Areolae are radiate and

coarsely punctate, with a polygonal outline. Areolae also become slightly larger

towards the valve center. Areolae open as ringed pores on the interior of the 68

valve. Labiate processes are visible most commonly under SEM. Pseudonodule

is obscure in LM, and most commonly observed under SEM. Mantle areolae

are closely packed, and often break to create a “jagged” appearance in imperfectly

preserved specimens.

Type Locality: Beurn (Hessen), Fossil, Germany.

Remarks: This species is most easily discernable from A. motilis by the significantly

flatter valves, coarser punctae, and the lack of a variable hyaline area in the center

of the valve. Krasske (1934) originally made remarks associating this species

with the genera Fragilaria and Aulacoseira; these associations hold true in these

samples.

Distribution: This species is very rare, being seen in the lower section of the Top Crude

unit, and most commonly seen in the Frenchman Hills and near the Gorge

Amphitheater. Occurrences are rare in the ALP.

Actinocyclus motilis Bradbury and Krebs 1995

Plate 17 number 1-5, Plate 18 number 5-6

Basionym: Actinocyclus motilis Bradbury and Krebs 1995

Synonyms: Cestodiscus mobilis Lohman 1954, Coscinodiscus subtilis Ehrenberg 1841

sensu Vanlandingham 1964

Description: Circular, concentrically undulate valves that become more flat in smaller

specimens. Diameter ranges 25 – 80m. Areolae are polygonal with round

openings on valve interiors, and are larger in the center of the valve. Labiate

processes are often visible in LM, and may be long stemmed. Pseudonodule most 69

often seen in SEM and is obscure in LM. Hyaline area in the center is variable,

and best developed in larger specimens. Areolae in the center may be loosely

packed. Short hyaline rows caused by the stellate pattern of areolae row

termination on the valve mantle are visible in LM, though are often overlooked

and may be best noticed on close examination of light micrographs.

Type Locality: Esmerelda Fm, Cedar Mountain, Mineral County, Nevada, USA.

Remarks: This species has a wide size variation and is by far the most common

Actinocyclus species in the deposit. As with A. krasskei, this species is highly

susceptible to breakage and fragments are common across all units. Bradbury and

Krebs (1995a) mention this species as being similar to the illustrations of

Coscinodiscus subtilis Ehrenberg 1841 in Vanlandingham (1964), but they are

ambiguous if they thought the species concept extended to this material. All

designations of Coscinodiscus subtilis and C. near subtilis by Vanlandingham

(1964) are considered A. motilis here.

Kocioleck and Spaulding (2001) interpreted an Actinocyclus sp. as being

morphologically variable with E. baileyi (listed incorrectly as E. arenaria). The

Actinocyclus they illustrate is likely A. motilis (Fig 7, b and d). Rare occurrences

of A. motilis and E. baileyi valves fitted together are seen in the Quincy

Diatomite. These rare occurrences are most commonly seen in the lower section

of the Top Crude where Actinocyclus species dominate.

Distribution: Observed as fragments in all samples, though it most commonly occurs in

the lower portions of the Top Crude, and in the Four Crude, where the genus 70

Actinocyclus is most common. It is observed in the Frenchman Hills, Gorge

Amphitheater, and the ALP.

Actinocyclus sp.

Plate 18 number 4

Description: Valves circular with concentric undulation. Diameter ranges at least 35–

55m. Areolae are radiate and made of porous clusters that open as slit-like pores

in the valve interior. Labiate processes are at least 1μm long, with several being

visible along the margins of the valve.

Remarks: This species is only recognizable under SEM, making it very rare. It is figured

in Kocioleck and Spaulding (2002), fig. 5, d and f. It is left in open nomenclature

pending further observations and character description.

Distribution: This species has only been observed in the ALP.

Order: Paraliales Crawford 1990

Family: Paraliaceae Crawford 1988

Genus: Ellerbeckia Crawford 1988

Ellerbeckia baileyi Crawford 2007

Plate 3 number 1-5, Plate 21 number 4-5

Basionym: Cestodiscus baileyi Smith 1877

Synonyms: Melosira baileyi (Smith) Wolle 1890, Coscinodiscus baileyi (Smith) Rattray

1890, Melosira sulcata (Ehrenberg) Kützing sensu Vanlandingham 1964,

Ellerbeckia arenaria (Moore) Crawford sensu Kociolek and Spaulding 2002 71

Description: Flat, large diameter valves with diameter ranging 32 – 70μm. Valves

are almost always in girdle view due to the large height to diameter ratio (>1:3).

Valve margin of specimens is densely poroid, with a large hyaline area in the

center. Linking grooves are visible on the valve surface, and are usually

arranged on the interior of the valve, with rare occurrences of these grooves

extending to the valve margin. Groove depth is slightly variable, with those

specimens showing grooves restricted to the valve interior exhibiting the deepest

grooves. Mantle processes are visible under SEM, with several processes lining

the entire mantle of each valve. All specimens exhibit heavy silicification.

Type Locality: Lost River, lower Klamath Lake, OR, USA

Remarks: This is the only species of Ellerbeckia in the Quincy Diatomite making it very

distinctive among the flora. It is almost always well preserved with minimal

breakage. Dominance in the Bottom Crude is greatest at the base of the unit, and

becomes reduced near the top.

Kocioleck and Spaulding (2002) discuss the possibility of this species

(misidentified as E. arenaria) being an internal valve of Actinocyclus and that the

ability to produce Actinocyclus sp. valves was lost as Ellerbeckia began to inhabit

fresh water in the Miocene. The dominance of E. baileyi in the

Bottom Crude may provide a more constrained timing for transition away from

the hypothesized heterovalve condition and the incursion of Ellerbeckia into

fresh water systems.

72

Distribution: This species is found in all three sublayers, though it diagnoses the Bottom

Crude due to its abundance (at least 25%). It is most common in the Frenchman

Hills. This location in Grant County, WA is the third confirmed location for E.

baileyi (Crawford 2007).

Class:

Subclass: Fragilariophycidae

Order: Fragilariales Silva 1962

Family: Fragilariaceae Greville 1833

Genus: Diatoma Bory de Saint-Vincent 1824

Diatoma anceps (Ehrenberg) Kirchner 1878

Plate 11 number 8

Basionym: Odontidium anceps Ehrenberg 1841

Synonyms: Fragilaria ? anceps Ehrenberg 1843, Odontidium captiatum Rabenhorst

1853, Odontidium anomalum Smith 1856, Odontidium anceps (Ehrenberg)

Ralphs in Pritchard 1861, Odontidium anceps f. minor (Ehrenberg) Rabenhorst

1864, Odontidium anceps f. intermedia (Ehrenberg) Rabenhorst 1864, Diatoma

anceps var capitata Peragallo in Tempère and Peragallo 1908, Diatoma anceps

var. constricta Tempère and Peragallo 1912.

Type Locality: Pelham, Massachusetts, USA.

Description: Valves are linear in outline with distinctly capitate to subcapitate ends.

Capitate ends are often as wide as the specimen’s maximum width. Length

ranges 13 – 25μm, and width ranges 3 – 6μm. Anomalous specimens may be 73

found that are significantly wider (up to 8m). Striae are very fine with numbers

variable between costae. Costae are not well developed, with 2 –6 costae per

10μm. Pseudoraphe is very narrow and obscure in LM.

Distribution: This species is rare across all samples.

Remarks: There appear to be two end members of morphology related to the main

body of the valve; some with length >> width, and those where length ~ width

(and often have subcapitate, protracted apices). Most specimens observed belong

to the length >> width group.

Diatoma hyemale (Roth) Heiberg 1863

Basionym: Fragilaria hiemalis Lyngbye 1819

Synonyms: Conferva hyemalis Roth 1800, Odontidium hyemale (Roth) Kützing 1844.

Description: Valves are elliptical to rhombic lanceolate in outline, with rare specimens

being orbicular. Length ranges 15 – 25μm and width ranges 14 – 20μm. Striae

are distinctly punctate, with punctae being less dense directly over costae, with 5

– 9 striae between costae. Costae are heavily silicified, with 3 – 4 costae per

10μm. Pseudoraphe is narrow but distinct in LM; it is rectangular in outline

Type Locality: “In piscins prope Schoenebeck non procul vegesack primo obseravi,

postea passim in fossis.”

Remarks: This species is most commonly seen in the lanceolate shape, at approximately

15m long. It might superficially be confused with Tetracyclus lanceolata, but

is far too small and costae are less densely packed per 10m. In its rare orbicular forms it may also be confused with T. elliptica, but is again, significantly smaller. 74

Vanlandingham (1964) provides a larger size range, however it is thought he

confused these with species of Tetracyclus. All specimens here are restricted to

the small size range.

Distribution: Rare in all samples. It occurs most commonly in the ALP in the upper-mid

portions of the Top Crude.

Diatoma hyemale f. mesodon (Kützing) Forti 1899

Plate 12 number 8-11

Basionym: Fragilaria mesodon Ehrenberg 1840

Synonyms: Odontidium mesodon Kützing 1844, Odontidium heimale var. mesodon

(Ehrenberg) Grunow 1862.

Description: Valves are elliptical to rhomboid elliptical in outline. Length ranges 8 –

25μm, and width ranges 6 – 13μm. Striae are distinctly punctate, with the

spacing between punctae becoming sporadic closer to the pseudoraphe; 8 – 10

striae between costae and approximately 2 costae per 10μm. Pseudoraphe is

poorly developed but visible in LM, and is lanceolate in outline.

Type Locality: Schwarzenberg bei Sachsen.

Remarks: This species variant was most commonly seen in its smallest forms, being near

circular and very small (approximately 10m). It is distinguished from D.

hyemale proper by generally being more circular/elliptical and by being smaller.

Distribution: Rare in all samples. It occurs most commonly in the ALP in the lower

portions of the Top Crude. It may be confused with Tetracyclus elliptica or T. 75

polygibbum, however it is smaller than both of those species and does not have a

well developed apical porefield.

Genus: Fragilaria Lyngbye 1819

Fragilaria lapponica Grunow 1881

Plate 15 number 7 & number 10

Basionym: Fragilaria lapponica Grunow 1881

Synonyms: none

Description: Valve outline is rhomboid elliptical to linear elliptical in larger specimens

and have broadly rounded apices. Length ranges 6 – 14μm and width ranges 3 –

4μm. Striae are very coarse and also appear inset into the valve surface, with

distinct relief visible between striae. Striae also are restricted to the valve

margin, with 8 – 10 striae per 10μm. Pseudoraphe area is very large due to short

striae and is rectangular in shape.

Type Locality: Uncertain

Remarks: This species is most easily distinguished from F. lenoblei by its more elliptical

valve outline, coarser striae, and significantly larger pseudoraphe central area.

There may also be some superficial confusion with F. martyi as it is similarly

striate; however F.lapponica is again more elliptical and has a larger

pseudoraphe.

Distribution: Found exclusively in the Frenchman Hills, in the Bottom Crude and Four

Crude units. This species is usually rare, with a few samples showing slightly

increased abundance. 76

Fragilaria near lenoblei Manguin 1952

Plate 15 number 11-12

Basionym: Fragilaria lenoblei Manguin 1952

Synonyms: none

Description: Valve outline is elliptical lanceolate with broadly rounded apices. Length

ranges 8 – 12μm and with a 2 – 3μm width range. Striae are marginal, coarse and

very short. Specimens display a large, lanceolate shaped pseudoraphe, with 10 –

12 striae per 10μm.

Type Locality: Antsirabe, Madagascar (Fossil).

Remarks: This species is distinct from F. lapponica based on the smaller striae and the

slightly different valve outline. Pseudoraphe shape is also more distinct in F.

lenoblei specimens than in other Fragilaria species.

Distribution: Very rare in the Frenchman Hills and absent in the Gorge Amphitheater and

ALP. It was only observed in the Bottom Crude unit.

Fragilaria near socia (Wallace) Lange-Bertalot 1981

Plate 15 number 9 & number 14

Basionym: Synedra socia Wallace 1960

Synonyms: none

Description: Valve linear lanceolate to elliptical lanceolate with sub-rostrate apices.

Length ranges 17 – 25μm and width ranges 3 – 4μm. Striae are parallel with 16 –

17 per 10μm. Pseudoraphe is narrow. Central area is devoid of striae and appears 77

as a hyaline area. A slight constriction then swollen section marks the boundaries

of this central area.

Type Locality: Savannah River, Screven Co., Georgia, USA.

Remarks: All measurements for specimens assigned to this designation fit the

description in Patrick and Riemer v. I (1966) with illustrations for Fragilaria

socia being an excellent match. However, I am unable to find this species

previously reported in the fossil record. The species matching this description are

therefore called F. near socia.

Distribution: Very rare across all sampling locations, though it is most commonly found

in the Frenchman Hills in the Bottom Crude unit. It also appears in the upper

most sections of the ALP Top Crude.

Fragilaria vaucheriae (Kützing) Peterson 1938

Plate 21 number 11-12

Basionym: Exilaria vaucheriae Kützing 1833

Synonyms: Synedra vaucheriae (Kützing) Kützing 1844, Synedra vaucheriae var.

parvula (Kützing) Rabenhorst 1864, Synedra vaucheriae var. distans Grunow in

Van Heurk 1881, Fragilaria intermedia Grunow in Van Heurk 1881.

Description: Valves lanceolate in outline with acutely rounded apices. Some specimens

are “crooked”, displaying a clear bend and offset of the valve outline in the

central area. Length ranges 10 – 18μm and width ranges 2 – 4μm. Striae are

straight, even across samples with the crooked center, with 15 – 16 striae per

10μm. Central area of valve has one side where striae are interrupted and 78

discontinuous, creating a hyaline area on one side of the valve. This hyaline area

is of variable size between specimens. Pseudoraphe is narrow.

Type Locality: On Vaucheria clavata in einer Quelle bei Weissenfels.

Remarks: Most specimens were observed with the “crooked” center. Some smaller

specimens had such large hyaline central areas where the continuous striae were

difficult to see.

Distribution Very rare across the Frenchman Hills, it was not encountered in the Gorge

Amphitheater, but was noticed once in the ALP.

Genus: Fragilariforma Williams and Round 1987

Fragilariforma virescens (Ralphs) Williams and Round 1987

Plate 15 number 15-19

Basionym: Fragilaria pectinalis Ehrenberg 1832

Synonyms: Diatoma sulphurascensi Agardh 1832, Fragilaria virescens Ralphs 1843,

Diatoma virescens Hassall 1845, Fragilaria aequalis Heiberg 1863,

Fragilaria virescens var. bohemia Grunow 1881, Fragilaria virescens var.

producta (Lagerstedt) De Toni 1891,Fragilaria virescens f. elongata Héribaud

1893, Nematoplata virescens Kurtze 1898, Fragilaria aequalis var. major

Tempère and Peragallo 1910, Fragilaria virescens Patrick and Reimer 1966.

Description: Valves are linear with protracted rostrate apices. Length ranges 15 – 43μm

and width ranges 4 – 5μm. Striae are linear, with individual punctae varying

between clearly visible to obscure in LM; striae range from 15 – 18 per 10μm. 79

Rare, smaller specimens may be found with striae slightly offset from the

opposite side of the valve. Pseudoraphe is clear but very narrow.

Type Locality: In freshwater pools. Cold Bath, Tunbridge Wells, leg. Madron and

Chyanhl Moor, near Penzance, England.

Remarks: The long, linear nature of this species makes it very distinct among the

Fragilaria of the Quincy Diatomite.

Distribution: Found rarely in the Frenchman Hills in all three units and rarely in the

Gorge Amphitheater, it is most commonly encountered in the ALP, where it is the

most abundant Fragilaria species through all samples at that locality.

Fragilariforma intortus nom. nov.

Plate 22 number 1-10, Plate 23 number 1-8

Description: Broadly lanceolate, with rostrate to capitate apices. Length appears ranges

from 25 – 50μm, with width ranging 8 – 9μm. Striae are uniseriate and punctate,

though individual punctae may be obscure in less well preserved specimens.

Striae are straight, with heavily obscured pseudoraphe; 9 – 10 striae per 10μm.

Only the largest specimens show a break in the striae producing a true

pseudoraphe, most easily seen in the largest striae in the center of the frustule.

Specimens are asymmetrical about the transapical axis, producing a rotational

symmetry; expression of this asymmetry is variable between specimens, with

some specimens appearing nearly straight, and others being strongly asymmetric.

Large apical pore fields are visible in LM and SEM, with distal striae sometimes 80

being distorted and not straight. Spines present along valve margin at base of

striae.

Type Location: Quincy Diatomite, Frenchman Hills, Grant Co, WA. USA.

Distribution: This species is equally distributed across all localities and units. It is

most common in the Frenchman Hills in the Bottom Crude

Remarks: It appears Vanlandingham was simply unsure of what to call this species;

Vanlandingham 1964 (plt. 27, fig. 26, 27) called this Fragilariopsis (?), and

showed more straight specimens with slightly less distinct interruptions between

the striae. In Vanlandingham 1967 (plt. 11, 20, 21) the same as the species

identified as Synedra (?), with those specimens being very similar except for a

small difference in length and the slightly more visible asymmetry about the

transapical axis. This asymmetry, however, is visible in even the straightest

looking specimens and is a diagnostic feature.

This species is assigned to Fragilariforma based on the description of F.

virescens by Williams (2001) and Williams and Round (1987), where the

presence of rostrate to capitate apices, a simple apical pore field, and the presence

of a single, polar rimoportulae, and the presence of spines behind the striae are

used as the basis of the genus. Species name is based on the Latin word for

crooked.

Genus: Staurosirella Williams and Round 1988

Staurosirella martyi (Héribaud) Morales and Manoylov 2006

Basionym: Opephora martyi Héribaud 1902 81

Synonyms: Fragilaria mutabilis f. martyi (Héribaud) Cleve-Euler 1932, Martyana martyi

(Héribaud) Round 1990, Fragilaria leptostauron var martyi (Héribaud) Lange-

Bertalot 1991, Fragilaria martyi (Héribaud) Lange-Bertalot 1993.

Description: Valve outline is slightly variable; most specimens are generally elliptical

lanceolate, though rare cuneate or clavate shaped specimens can be found.

Asymmetrical about the transapical axis. Apices are broadly rounded. Length

ranges 6 – 15μm with width ranging 3 – 7μm. Striae are lineate and straight,

possibly becoming slightly radiate near the apices with 6 – 10 striae per 10μm.

Pseudoraphe is narrow and lanceolate in shape.

Type Locality: Cantal, Neussargues, France.

Remarks: The narrow pseudoraphe and elliptical lanceolate valve outline are the most

diagnostic features of this species compared to other Fragilaria found in the

deposit.

Distribution: Very rare in the Frenchman Hills, being found only in the Bottom Crude

unit. It is absent from the ALP.

Staurosirella leptostauron (Ehrenberg) Williams and Round 1987

Basionym: Biblarium leptostauron Ehrenberg 1854

Synonyms: Staurosira pinnata Ehrenberg 1854, Odontidium ? harrisonii Smith 1856,

Fragilaria harrisonii (Smith) Grunow 1862, Fragilaria leptostauron (Ehrenberg)

Hustedt 1931.

Description: Valve outline is broadly rhombic and inflated to nearly cuneate in some

specimens; all specimens have rostrate apices. Length ranges 11 – 23μm and 82

width ranges 7 – 14μm. Striae are singly punctate with the punctae being distinct

under LM, and appearing granulate in some samples; 6 – 9 striae per 10μm.

Pseudoraphe is narrow and clearly extends the length of the valve in LM.

Type Locality: “Torf von Newhaven, Connecticut, Nord-Amerika, T.V. ii, F. 24.”

Remarks: This species is unique among the Fragilariforms found in this sample as it is so

strongly punctate, making it one of the most easily identified species in the

deposit.

Distribution: Very rare, it is restricted to the Frenchman Hills in the Bottom Crude unit. It

is absent from all other samples and localities.

Staurosirella pinnata var. lancettula (Schumann) Siver annd Hamilton 2005

Plate 15 number 8

Basionym: Fragilaria lancettula Schumann 1867

Synonyms: Fragilaria pinnata var. lancettula (Schumann) Hustedt 1913

Description: Valve outline is rhomboid elliptical with broadly rounded apices. Length

ranges 10 – 20μm and width ranges 3 – 6μm. Striae are straight and singly

punctate with 10 – 12 striae per 10μm. Punctae of striae are clearly visible in

LM. Pseudoraphe is very narrow and may be slightly obscure in LM with smaller

specimens.

Type Locality: Russia. Kaliningrad Oblast.

Remarks: This species is superficially similar to F. leptostauron, however it is not as

wide and has more striae per 10μm. 83

Distribution: This species is rare across all three sampling localities and was usually

encountered as solitary specimens on a slide. It is found in all three units in the

Frenchman Hills, with greatest abundance being in the Four Crude unit.

Genus: Meridion Agardh 1824

Meridion circulare (Greville) Agardh 1830

Plate 10 number 6, Plate 19 number 3 & number 7

Basionym: Echinella circularis Greville 1822

Synonyms: Meridion zinkenii Kützing 1843, Meridion circulare var. zinkenii (Kützing)

Grunow 1862.

Description: Valves are claviate to linear, with one apex being broadly rounded and the

other protracted. Length ranges 20 – 39μm, with width ranging 5 – 8μm. Costae

are poorly preserved and obscure in LM, appearing as irregular gaps between the

striae. Striae occur between the costae. Striae at straight in the central portions of

the valve and radiate at the ends of the apices. Striae are uniseriate with

occasional isolated punctae occurring along the narrow margin of the

pseudoraphe. Striae are also offset across the pseudoraphe; 10 – 14 striae per

10μm. Pseudoraphe is very narrow and clearly visible along the length of the

valve.

Type Locality: Rivulet near Dumbryden Quarries

Remarks: Patrick and Reimer v. 1 (1966) list this species as having more striae per 10μm

(15 – 16), but the assignment appears correct with form and other references on

Miocene taxa. 84

Distribution: Rare in all samples where it is encountered.

Genus: Pseudostaurosira Williams and Round 1987

Pseudostaruosira brevistriata (Grunow) Williams and Round 1987

Plate 15 number 1

Basionym: Fragilaria brevistriata var. subacuta and var. pusilla Grunow 1881

Synonyms: Fragilaria brevistriata Grunow 1885

Description: Valves are linear lanceolate to rhombic lanceolate with rostrate apices.

Some specimens have slightly constricted mid-valve regions. Length ranges 15 –

24μm, and width ranges 3 – 5μm. Striae are parallel, singly punctate and very

short, creating a broad central area for the pseudoraphe; 14 – 16 striae per 10μm.

Pseudoraphe is broad and lanceolate in appearance.

Type Locality: Fresh water, Bruxelles (Delogne).

Remarks: This species shares a similar outline with larger specimens of S. construens v.

venter; however the large, lanceolate shaped pseudoraphe is the most diagnostic

features of this species.

Distribution: Very rare is the Frenchman Hills and Gorge Amphitheater, and absent from

the ALP. It is most commonly found in the Bottom Crude unit.

Pseudostaurosira brevistriata var. elliptica (Héribaud) Kingston 2000

Basionym: Fragilaria brevistriata var. elliptica Héribaud 1903

Synonyms: none 85

Description: Valve outline is elliptical lanceolate. Length ranges 6 – 8μm, and width

ranges 3 – 4μm. Striae are singly punctate with elliptical/rectangular punctae that

are clearly visible in LM, with 13 – 14 striae per 10μm. Broad, lanceolate

pseudoraphe area is visible in LM.

Type Locality: Cantal, dépot de Moissac, France.

Remarks: Most easily distinguishable from P. brevistriata by having the elliptical

lanceolate outline without protracted apices.

Distribution: This species is very rare, and was only encountered twice in the Frenchman

Hills. It is absent from the Gorge Amphitheater and the ALP.

Pseudostaurosira brevistriata var. subcapitata (Grunow) nom. nov.

Plate 15 number 2-3, Plate 21

Basionym: Fragilaria brevistriata var. subcapitata Grunow 1881

Van Heurck, Syn. Diat. Belg. pl. XLV: fig. 33. 1881

Synonyms: none

Description: Valves have linear sides with subcapitate apices. Striae are short, but often

longer than the nominal variety. Length of this variety ranges 15 – 27μm, and

width ranges 3 – 5μm. Approximately 14 striae per 10μm. Large, lanceolata

shaped pseudoraphe.

Type Locality: “Bruxelles (Delogne)”

Remarks: This species is most easily distinguished from the longest valves of Staurosira

construens var. venter by a much larger pseudoraphe and the protracted, 86

subcapitate apices. It is distinguished from P. brevistriata proper by the distinctly

subcapitate ends and linear sides of the valve.

The nominal variety of this species, P. brevistriata was transferred out of the genus

Fragilaria by Williams and Round (1988); however this variety has not yet been

transferred. Its transfer to Pseudostaurosira is proposed here.

Distribution: Very rare in the Frenchman Hills, only occurring in the Bottom Crude and

uppermost portions of the Four Crude units. Absent from the Gorge

Amphitheater and ALP.

Genus: Staurosira (Ehrenberg) Williams and Round 1987

Staurosira construens (Ehrenberg) Williams and Round 1987

Plate 15 number 4-5

Basionym: Staurosira construens Ehrenberg 1843

Synonyms: Odontidium tabellaria Smith 1856, Dimerogramma tabellaria Ralphs in

Pritchard 1861, Fragilaria construens (Ehrenberg) Grunow 1862.

Description: Valves have an inflated central area and protracted apices. Length ranges 11

– 14μm and width ranges 5 – 8μm. Striae are parallel, and lineate with

elliptical/rectancular pores; 13 – 15 striae per 10μm. Pseudoraphe is very narrow,

uniform in width, and clearly visible in LM.

Type Locality: “…Newhaven in Nord-Amerika…”

Remarks: S. construens always has an inflated central portion with protracted apices in

this material, making it easily differentiable from S. construens var. venter. The

more lanceolate shaped valves of this species are absent in all samples. 87

Distribution: Rare in all samples from the Frenchman Hills. It is absent from the ALP.

Staurosira construens var. venter (Ehrenberg) Hamilton 1992

Plate 16 number 1-16

Basionym: Fragilaria venter Ehrenberg 1859

Synonyms: Fragilaria construens var. venter (Ehrenberg) Grunow 1881

Description: Valves are elliptical lanceolate to linear lanceolate, though smaller

specimens are nearly circular and longer specimens may have slight constrictions

in the center. Length ranges 4 – 21μm and width ranges 3 – 7μm. Striae are

straight, extend close to the valve center with a very narrow pseudoraphe. Close

examination may be necessary to make this determination as poorly developed

striae may be hard to see in LM; 13 – 16 striae per 10μm.

Type Locality: Various localities in Germany, France, Peru, Bohemia, and Hungary

Remarks: Wide variation in size is seen in this species. It is most commonly encountered

in the 10 – 18μm lengths. Striae may be poorly developed, creating superficial

confusions with Pseudostaurosira brevistriata var. subcapitata in larger

specimens. The narrow pseudoraphe is the best distinguishing factor for this

species.

Thomas (1981) discusses this species as being most abundant in samples with

significant silt deposits, possibly being an ecologic indicator of high runoff areas.

Due to the purity of the Quincy Diatomite and the great abundance of this

species, this speculation is interpreted as incorrect. 88

Distribution: Occurs across all sampling localities and is observed in most samples. It is

most abundant in the Bottom Crude unit in the Frenchman Hills, and

progressively rarer moving north to the Gorge Amphitheater and the ALP. This is

by far the most abundant Fragilaria species in the Quincy Diatomite.

Genus: Synedra Ehrenberg 1832

Synedra tabulata var. fasciculata (Kützing) Hustedt 1932

Basionym Synedra fasiculata Kützing 1844

Synonyms: Synedra (afrinis var.) fasiculata (Kützing) Grunow in Van Heurk 1881.

Description: Valves are linear, with protracted rostrate apices. Length ranges 30 – 45μm

and width ranges 5 – 6μm. Striae are lineate and straight throughout the valve; 11

– 15 striae per 10μm, with the central portion of the valve face being less densely

striate. Pseudoraphe is lanceolate in shape, and is very large, accounting for

approximately half the valve face.

Type Locality: Uncertain

Remarks: The length here is significantly smaller than those presented by Schauderna

(1983), who lists the length range as 30 – 70μm. This species may need to be

transferred to Fragilaria as several other varieties have already been transferred

(Lange-Bertalot 1981), and the features of this valve appear more closely allied

with Fragilaria. However, as only three specimens were encountered, it is left in

its current taxonomic position pending further observations.

Distribution: This species is very rare. It is restricted to the Frenchman Hills, and is only

found in the Bottom Crude and Four Crude units. 89

Order: Tabellariales Round 1990

Family: Tabellariaceae Kützing 1844

Genus: Tetracyclus Ralfs 1843

Tetracyclus ellipticus (Ehrenberg) Grunow 1862

Plate 9 number 1

Basionym: Biblarium ellipticum Ehrenberg 1845

Synonyms: Biblarium compressum Ehrenberg 1845, Tetracyclus compresus (Ehrenberg)

Peragallo in Héribaud 1893, Tetracyclus ellipticus f. minor Hustedt in Schmidt

1912

Description: Valves elliptical to circular in outline with broadly rounded apices. Apical

pore fields are present. Length ranges 30 – 42μm and width ranges 25 – 40μm

wide. Costae are very robust, and are usually primary, rarely secondary. Striae

are parallel and distinctly punctate, with 8 – 10 striae in between costae. Density

remains relatively constant directly over costae as in between. Pseudoraphe is

very narrow and has a slight increase in silicification (sternum) on the interior of

the valve. Several rimoportulae are visible under SEM.

Type Locality: “Sibiria. Oregon. Fossile.” Columbia River, OR.

Remarks: This species is superficially similar to T. polygibbum and Diatoma

hyemale, however it is larger than Diatoma hyemale, and is more elliptical and

circular than T. polygibbum. The presence of apical pore fields is also

diagnostic.

Distribution: This species is very rare across all sampling localities. It is most commonly

found in the Frenchman Hills in the Top Crude. 90

Tetracyclus lacustris Ralfs 1843

Plate 9 number 3

Basionym: Tetracyclus lacustris Ralfs 1843

Synonyms: Tetracyclus stella (Ehrenberg) Héribaud 1902

Description: Valves are triundulate, creating the appearance of a tri–section valve.

Undulations in valve outline vary between nearly equal, to a greatly exaggerated

central undulation and highly reduced apical undulations. Central area is slightly

swollen compared to the rest of the valve. Length ranges 20 – 45μm, with width

ranging 7 – 15μm. Costae are strongly developed, with 2 – 3 per 10μm. Striae

are straight and distinctly punctate, though punctae are often discontinuous across

single striae, particularly near the valve face center; 7 – 10 striae in between

costae. Pseudoraphe is narrow with a moderately developed sternum on the valve

interior. 1 rimoportulae is visible under both SEM and LM, and is usually

close to the pseudoraphe on the valve face.

Type Locality: Llyn Prefeddyr near Barmouth, Rev. T. Salwey; pool near Dolgelley,

Wales.

Remarks: Though rare, the distinctive valve outline makes this species the most readily

identifiable Tetracyclus in the Quincy Diatomite.

Distribution: This species is rare across all sampling localities. It is most common in the

Frenchman Hills in the sections of the Top Crude unit.

91

Tetracyclus lancea (Ehrenberg) Peragallo in Héribaud 1893

Plate 20 number 1

Basionym: Biblarium lancea Ehrenberg 1845

Synonyms: Tetracyclus ellipticus var. lancea f. elongata Hustedt in Schmidt 1912

Description: Valves are lanceolate. Length ranges 65 – 100μm and width ranges 15 –

25μm. Length to width ratio is always greater than 4:1. Costae are strongly

silicified, usually primary, with 3 – 4 per 10μm. Striae are straight with 5 – 7

between costae. Pseudoraphe is very narrow with sternum on the interior of the

valve. 1 – 2 rimoportulae are sometimes visible in LM, but most commonly seen

under SEM.

Type Locality: “In Oregonia Fossile.”

Remarks: This species is very large (possibly the largest species in the Quincy

Diatomite). It is easily distinguished based on its size and strongly lanceolate

shape.

Distribution: This species is very rare across all sampling localities. It is uniformly

distributed across the deposit without any particular unit or sampling locality of

greater abundance.

Tetracyclus lata (Hustedt) Williams 1996

Plate 10 number 3

Basionym: Tetracyclus ellipticus var. lancea f. lata Hustedt in Schmidt 1912

Synonyms: Tetracyclus ellipticus var constrictus Hustedt in Schmidt 1912 92

Description: Valves linear lanceolate to lanceolate. Length ranges 40 – 70μm and width

ranges 5 – 25μm. Costae are strongly developed, and usually primary, though

may be secondary; 2 – 4 costae per 10μm. Striae are straight, with 8 – 10

between costae. Punctae of striae are usually distinct in LM. Pseudoraphe is

narrow, but the accompanying sternum is obscure in LM and is likely

underdeveloped in these specimens. A single rimoportulae is visible under SEM

on the valve face.

Type Locality: “Briete Busch bie Hainspach.”

Remarks: Some specimens are more lanceolate in outline and may resemble T. lancea,

however T. lata is smaller and not as wide. Williams (1996) lists 1 – 3

rimoportulae being possible on the valve face or mantle, however only specimens

with a single rimoportulae on the valve face were observed.

Distribution: Rare across all sampling localities and units, this is, however, the most

common species of Tetracyclus found in the Quincy Diatomite. It is found most

frequently in the Frenchman Hills in the Four Crude unit.

Tetracyclus linearis (Ehrenberg) Grunow 1862

Plate 9 number 2

Basionym: Biblarium lineare Ehrenberg 1845

Synonyms: Biblarium lamina Ehrenberg 1845, Tetracyclus lamina (Ehrenberg) Héribaud

1893, Tetracyclus ellipticus var. lamina (Ehrenberg) Peragallo 1903, Tetracyclus

lamina f. lata Tempère et. Peragallo 1909, Tetracyclus ellipticus var. linearis 93

(Ehrenberg) Hustedt in Schmidt 1912, Tetracyclus ellipticus var. lamina f. lata

(Tempère et. Peragallo) Mills 1935.

Description: Valves are linear lanceolate. Length ranges 40 – 75μm and width ranges 20

– 30μm. Costae are robust and almost always primary, with 2 – 3 per 10μm.

Striae are straight and clearly punctate, with 5 – 8 between costae. Pseudoraphe is

narrow with a poorly developed sternum on the valve interior. A single

rimoportulae is visible under SEM, located on the valve face close to the margin.

Type Locality: “Fossil ad Bargus. Sibiriae et Oregonia.”

Remarks: Williams (1996) lists this species minimum length as being 50μm. Smaller

species have been observed here, and the minimum length is thus reduced to

40μm.

Distribution: This species is rare across all sampling localities and units. It is most

common in the ALP near the bottom of the exposed section of Top Crude.

Tetracyclus polygibbum (Pantocsek) Jousé 1952

Plate 10 number 1-2

Basionym: Stylobiblium polygibbum Pantocsek 1892

Synonyms: Stylobiblium ovale Pantocsek 1892, Stylobiblium haradaae Pantocsek 1892,

Tetracyclus jimboi Jousé 1952, Tetracyclus rhombus var producta Tempère et.

Peragallo 1909.

Description: Valves are elliptical to rhomboid elliptical with broadly rounded apices.

Length ranges 20 - 40μm, with a width ranging 7 – 10μm. Costae are strongly

developed, and may be primary, secondary, or tertiary; 1 – 3 costae per 10μm. 94

Striae are straight and clearly punctate, with 8 – 10 striae in between costae.

Striae density also appears slightly reduced directly over costae in some

specimens. Pseudoraphe is narrow and is clearly visible in LM; sternum is poorly

developed on the valve interior. 1 – 3 rimoportulae are located near the valve

margin, and are always near the center of the valve.

Type Locality: “In strasis teriariis aquae dulcis ad Sentenai in insula Jesso.”

Remarks: This species has previously only reported as T. sp. aff. polygibbum in North

America (Williams 1996). This is the first confirmed reporting from the NW

United States.

Distribution: This species is very rare and uniformly distributed across all sampling

localities and all units.

Tetracyclus williamsensii (Hustedt) nom. nov.

Plate 8 number 1-6, Plate 20 number 2-5

Basionym: Tetracyclus ellipticus var. lancea f. subrostrata Hustedt 1912.

A. Schmidt’s Atlas: pl. 281 fig. 17 and 18.

Description: Valves are linear lanceolate to rhombic lanceolate with rostrate to

subrostrate apices. Porefield is visible under SEM, and it is located on the valve

mantle and the tips of the apices. Length ranges 18 – 55μm, and width ranges 10

– 25μm. Costae are robust and almost always primary, rarely secondary; 3 – 4

transapical ribs per 10μm. Costae are straight in the central portion of the valve,

becoming concave towards the valve apices. Striae are more or less straight at the

valve center, and become radiate towards the apices. Striae are also 95

discontinuously punctate in some samples; 6 – 10 striae in between costae.

Pseudoraphe is very narrow, and is not perfectly straight across the valve.

Sternum is well developed and strongly silicified on the valve interior. 1

rimoportulae is visible under SEM; it most often located in between the central

costae, and is on the valve face near the mantle or rarely on the mantle. Septum

has a small concavity seen in girdle bands of this species. Girdle bands are

distinctive, with filamentous ends and an “ox-bow” shape not seen in other

Tetracyclus species.

Type Locality: Columbia River, Fossil.

Remarks: This species was figured in Schmidt’s Atlas by Hustedt as T. ellipticus var.

lancea f. subrostrata. The only other mention of this species found is by

Vanlandingham 1964 (pl. 48, 1 – 3). This species is too small to be part of the T.

lancea species concept as defined by Williams 1996, and its common form as 20

– 30μm long with distinctly subrostrate ends make it distinctive among the flora

of the Quincy Diatomite. The presence of 1 rimoportulae consistently near the

valve margin in between the central transapical ribs is also a distinguishing

feature from T. lancea. However, the forma name cannot be elevated to the

species level, as T. subrostrata was used to designate a species from a Chinese

Miocene fossil deposit (Williams 2008). The name T. williamsensii is therefore

used to recognize Dr. Williams contribution to the and description of

the genus Tetracyclus.

The source of Hustedt’s original fossil specimens collected from the Columbia

River is most likely the Quincy Diatomite because this unit is pervasive 96

throughout the Quincy Basin and is sufficiently unconsolidated to be a ready

source of reworked fossil material.

Distribution: Uniformly distributed between all sampling localities and units. It is most

common in the Frenchman Hills in the Bottom Crude.

Class: Bacillariophyceae

Subclass: Eunotiophycidae

Order: Eunotiales Silva 1962

Family: Eunotiaceae Kützing 1844

Genus: Eunotia Ehrenberg 1837

Eunotia curvata (Kützing) Lagerst 1884

Plate 11 number 9

Basionym: Synedra lunaris Ehrenberg 1832

Synonyms: Exilara lunaris Hassall 1844, Eunotia lunaris (Ehrenberg) Grunow 1877,

Pseudoeunotia lunaris (Ehrenberg) De Toni 1891, Synedra falcate Brébisson in

Kützing 1949.

Description: Valves are lunate to arcuate in shape with some specimens nearly straight;

broadly rounded apices. Length ranges 35 – 80μm and width is 3 – 4μm. Striae

are straight and extend across the entire valve face, conforming to the curvature of

the valve to stay straight; 13 – 18 striae per 10μm. Terminal nodules of raphe are

small and obscure in LM. Raphe is indistinct.

Type Locality: Berlin, Germany. 97

Remarks: This species is very rare and is most noticed at lower magnifications due to the

long length.

Distribution: This species is evenly distributed across all sampling localities. It is most

commonly found in the Top Crude unit in the ALP.

Eunotia pectinalis (Kützing) Rabenhorst 1864

Plate 12 number 6

Basionym: Himantidium pectinale Kützing 1844

Synonyms: Conferva pectinalis Müller 1788 (questionable), Conferva pectinalis DillW,

Eunotia pectinalis f. elongata Van Heurck 1881, Eunotia pectinalis var. stricta

(Rabenhorst) Van Heurck 1881

Description: Valves are linear, straight, and slightly elongated with broadly rounded

apices. Length ranges 20 – 41μm, and width ranges 5 – 8μm. Striae are straight

and extend across the entire valve surface, with 5 – 12 striae per 10μm. Terminal

nodules of raphe are very distinct at valve apices. Raphe is obscure in LM.

Type Locality: “In scaturiginibus Germaniae et Daniae.” (Müller)

Remarks: This is the most abundant Eunotia species in the Quincy Diatomite.

Distribution: This species occurs across all sampling localities, though it is restricted to

the Top Crude unit. It is most abundant in the upper section of the ALP.

Eunotia pectinalis var. minor (Kützing) Mueller 1910

Plate 12 number 7

Basionym: Himantidium minus Kützing 1844 98

Synonyms: Himantidium veneris Kutzing 1844, Eunotia impressa Ehrenberg 1854,

Eunotia pectinalis var. minus (Kutzing) Rabenhorst 1864, Eunotia pectinalis var.

impressa Müller 1898, Eunotia pectinalis var. minor f. impressa (Ehrenberg)

Hustedt in Pasch 1930.

Description: Valve outline is broadly linear with dorsal side being convex. Larger

specimens show an undulation in the dorsal side near the terminal nodules. Valve

apices are broadly rounded. Length ranges 14 – 31μm and width ranges 4 – 6μm.

Striae are straight at the valve center, becoming slightly concave towards the

apices, with 14 – 15 striae per 10μm. Terminal nodule is distinct in larger

specimens and slightly obscure in smaller specimens. Raphe is obscure in LM.

Type Locality: “Unter Süsswasseralgen bei Jever: Koch!, Germany.”

Remarks: This species variety was observed to coocur with the nominate variety, E.

pectinalis.

Distribution: This species is very rare and was only encountered once in the Frenchman

Hills, in the Top Crude unit.

Eunotia veneris (Kützing) De Toni 1892

Plate 12 number 1-5

Basionym: Himantidium veneris Kützing 1844

Synonyms: Eunotia incisa Gregory 1854

Description: Valve outline shows one side linear and the other side strongly convex.

Apices are rounded and tapered. Length ranges 13 – 23μm and width ranges 3 –

4μm. Striae are uniseriate, and straight in the central portions of the valve, 99

becoming slightly radiate towards the apices; 14 – 17 striae per 10μm. Raphe

body is obscure with terminal ends being distinct on the linear side of the valve.

Type Locality: Insel Trinidad, im Asphaltsee “Tacarigua”: Krüger, (Sonder!).

Remarks: The measurements given here represent the range of observed specimens.

Vanlandingham (1964) neglects to give a full range for dimensions (saying only

“approximately 15μm long and 3.5μm wide); the specimens on the larger end of

the range provided are confidently assigned based on striae counts and valve

outline.

Distribution: This species is very rare, however it is found in all sampling localities and

all three units. It is most commonly found at the ALP in the upper portions of the

Top Crude unit.

Subclass: Bacillariophycidae

Order: Naviculales Bessey 1907

Suborder: Neidiineae Mann 1990

Family: Cavinulaceae Mann 1990

Genus: Cavinlua Mann & Stickle 1990

Cavinula pseudoscutiformis (Hustedt) Mann and Stickle 1990

Plate 14 number 8

Basionym: Navicula pseudoscutiformis Hustedt 1930

Synonyms: none

Description: Valves are elliptical to orbicular. Length ranges 9 – 11μm, with width

ranging 7 – 8μm. Striae are strongly radiate and distinctly punctate. Striae in the 100

central area may be of variable length, and cut short by longer striae converging

in the central area; 26 – 29 striae per 10μm. Axial area is very narrow with a

circular to elliptical central area. Raphe is straight with terminal raphe ends being

indistinct in LM.

Type Locality: Zerstreut im Grundschlamm hosteinischer Seen, z.B. Plussee. Germany

Remarks: The fine striations are most diagnostic when comparing this species to

Navicula scuteloides.

Distribution: This species is very rare, being restricted to the Frenchman Hills in the

Bottom Crude unit.

Family: Neidiaceae Mereschkowsky 1903

Genus: Neidium Pfitzer 1871

Neidium near iridis var. ampliatum (Ehrenberg) Cleve 1904

Plate 21 number 3

Basionym: Navicula ampliata Ehrenberg 1854

Synonyms: none

Description: Valves are linear lanceolate with elongated, subrostrate apices. Length

approximately 50μm and width ranges approximately 15μm. Striae are straight,

becoming slightly convergent near the apices, and distinctly punctate; 15 striae

per 10μm. Longitudinal band on the valve face is broad, with a narrower

secondary band on the margin. Longitudinal bands have at least two rows of

punctae. Axial area becomes slightly narrower towards the center. Central area is

oval shaped. Raphe is filamentous, with proximal ends of raphe being hooked in 101

opposite directions. Distal ends of raphe are bifurcated and split to each side of

the valve face.

Type Locality: Jeusekguhre von Down, Mourne Mountains. Irland.

Remarks: This species was only found once under SEM. The specimen closely matches

the description and illustrations of N. iridis var. ampliatum in Patrick and Reimer

v.1 (1966) while being only slightly smaller, but as the specimen is at an angle, it

could be one of the varieties.

Distribution: Appears to be restricted to the ALP Top Crude.

Suborder: Sellaphorineae Mann 1990

Family: Sellaphoraceae Mereschkowsky 1902

Genus: Sellaphora Mereschkowsky 1902

Sellaphora bacillum (Ehrenberg) Mann 1989

Plate 10 number 5

Basionym: Navicula bacillum Ehrenberg 1843

Synonyms: none

Description: Valves are linear with broadly rounded apices. Length ranges 37 – 45μ,

with width ranging 8 – 12μm. Striae are radiate and uniseriate, with indistinct

punctae, and specimens have 15 – 18 striae per 10μm. Axial area is narrow, with

axial ridges visible under LM. Raphe is straight with hooked terminal ends.

Type Locality: Uncertain

Remarks: All specimens observed in this deposit are in the narrow range of length and

reduced width provided. This reporting is the oldest confirmed occurrence of the 102

genus Sellaphora, and the oldest confirmed occurrence of the species S. bacillum

(Evans et al. 2008, Mann pers. communication).

Distribution: This species rare in the Frenchman Hills and the Gorge Amphitheater, and

is absent from the ALP; it is most common in the Frenchman Hills in the Bottom

Crude unit.

Family: Pinnulariaceae Mann 1990

Genus: Pinnularia Ehrenberg 1843

Pinnularia nodosa (Ehrenberg) Smith 1856

Plate 10 number 4

Basionym: Navicula nodosa Ehrenberg 1838

Synonyms: Pinnularia nodosa var. genuine Cleve-Euler 1955

Description: Valves are triundulate, with undulations being fairly equal. Apices are

subrostrate to subcapitate. Length ranges 35 – 48μm and width ranges 7 – 9μm.

Striae are lineate and at least 1μm wide. Striae are radiate in the central portion of

the valve face, and transition to convergent near the apices. Central area striae are

absent, creating a larger central axial area; 10 – 11 striae per 10μm.

Type Locality: Bei Berlin, Germany.

Remarks: The dimensions observed are smaller than those listed by Vanlandingham

(1964) for length (46 – 48 μm), and slightly smaller than those given by Patrick

and Reimer v.1 (1966) for width (9 – 18μm).

Distribution: This species is very rare in the Frenchman Hills and Gorge Amphitheater.

It is most common in the ALP and the Top Crude unit. 103

Pinnularia near major (Kützing) Rabenhorst 1853

Plate 13 number 1-2

Basionym: Frustulia major Kützing 1833

Synonyms: Navicula major (Kützing) Kützing 1844

Description: Valves are linear with broadly rounded apices. Length ranges 100 – 150μm

and width ranges 23 – 27μm. Striae are lineate and at least 1μm wide. Striae are

radiate in the central portion of the valve face, and transition to convergent near

the apices. Length of striae is fairly consistent across most of the valve face, but

is slightly shorter in the central portion of the valve; 6 – 7 striae per 10μm. Axial

area is very broad, being at least 1/3 the breadth of the valve face. Raphe is

filiform, with indistinct terminal fissures, but distinct central nodules.

Type Locality: Halle, Germany.

Remarks: The lower end measurements of length and width are bellow those listed by

Patrick and Reimer v.1 (1966), and the species is thus designated near P. major.

All other parts of the description match observed specimens.

Distribution: This species is very rare and was only counted twice, however it is

interpreted to be equally distributed across all sampling localities and units as it

was found in the Frenchman Hills and ALP.

Suborder: Diploneidineae Mann 1990

Family: Diploneidaceae Man 1990

Genus: Diploneis Ehrenberg ex Cleve 1894 104

Diploneis ovalis (Hilse) Cleve 1894

Plate 14 number 5 & number 9

Basionym: Pinnularia ovalis Hilse in Rabenhorst 1861

Synonyms: none

Description: Valves are rhomboid elliptical with broadly rounded apices. Length ranges

22 – 35μm and width ranges 9 – 20μm. Striae are uniseriate and slightly radiate

with areolae detail obscure in LM. Longitudinal canals on either side of the axial

area are visible, with areolae surficial to the canals appearing as part of the striae

along the valve surface. Axial area is narrow with the raphe being slightly

obscure in LM. Central area is oval shaped and relatively small compared to the

valve surface. Raphe is straight, with proximal ends being distinct and terminal

fissures unresolved in LM.

Type Locality: “Strehlen in Schesien”

Remarks: Specimens observed were smaller than that observed by Vanlandingham

(1967), being on average 10μm shorter. However, the diagnostic central area

and longitudinal canals makes the author confident in this designation.

Distribution: This species is very rare. It is found in the Frenchman Hills in the Bottom

Crude unit. It is completely absent in the Gorge Amphitheater and the ALP.

Suborder: Naviculineae Hendey 1937

Family: Kützing 1844

Genus: Navicula Bory 1822 105

Navicula acceptata Hustedt 1950

Basionym: Navicula acceptata Hustedt 1950

Synonyms: Navicula ignorata var. acceptata (Hustedt) Lange-Bertalot 1984

Description: Valves are lanceolate with broadly rounded apices. Length ranges 5 –

12μm, with width ranging 3 – 5μm. Striae are straight across the length of the

valve, with a slightly increase in density near the apices. Punctae are indistinct

in LM. Axial area is narrow, with a small, circular central area. Raphe is straight,

with the terminal ends being obscure in LM in all but the most well preserved

specimens. Proximal raphe ends show very slight swelling.

Type Locality: Oberohe, Lüneberger Heide, Germany. Interglacial

Distribution: This species is very rare in the Quincy Diatomite. It is found at all sampling

localities, and is most abundant in the Top Crude unit.

Navicula reimerites Vanlandingham 1964

Plate 13 number 5-6

Basionym: Navicula reimertes Vanlandingham 1964

Synonyms: none

Description: Valves are lanceolate with broadly rounded apices. Apices have small

apical pore fields. Length ranges 11 – 23μm, with width ranging 3 – 6μm. Striae

are straight in the central portions of the valve, transitioning to slightly radiate

near the apices. Central portion of the valve is rectangular, with shortened striae

creating the larger area; 20 – 26 striae per 10μm. Axial is fairly broad, becoming 106

narrower towards the apices. Raphe is filamentous, with hooked terminal

fissures.

Type Locality: Quincy Diatomite, Grant County, Washington. USA. Fossil.

Remarks: The type locality given by Vanlandingham is in Townships and Ranges, and

corresponds exactly to the Gorge Amphitheater sampling locality (Mining section

20). The type locality listed here thus reflects the Quincy Diatomite being the

type deposit for this species.

Distribution: This species is rare, but present in all sampling localities. It is observed in

all three units; however it is found most commonly in the ALP in the upper

portions of the Top Crude unit. Vanlandingham (1964, 1967) reports it is also

found in the Savannah River, Georgia.

Navicula scutelloides Smith 1856

Plate 13 number 7

Basionym: Navicula scutelloides Smith ex Greg 1856

Synonyms: none

Type Locality: Fresh water, Ormsby, Norfolk (Mr. Bridgman); Rosthern Mere, Cheshire

(Dr. Arnott).

Description: Valves are elliptical to orbicular. Length ranges 12 – 14μm and width

ranges 10 – 12μm. Striae are of alternating short and long lengths, are distinctly

punctate, and are strongly radiate, with 10 – 13 striae per 10μm. Axial area is

very broad. Raphe is straight with indistinct terminal fissures. 107

Remarks: The clearly punctate striae and large axial area are the most diagnostic features

of this species.

Distribution: This species is very rare. It has been observed in the ALP, and was only

counted once in the Frenchman Hills in the Bottom Crude unit.

Order: Cymbellales Mann 1990

Family: Cymbellaceae Greville 1833

Genus: Cymbella Agardh 1830

Cymbella cistula (Hemprich and Ehrenberg) Kirchner 1878

Plate 14 number 1-4 &number 10

Basionym: Cocconema cistula Hemprich and Ehrenberg 1832

Synonyms: Bacillaria cistula Hemprich and Ehrenberg 1828

Type Locality: Uncertain

Description: Valves are cymbelloid with broadly rounded to slightly reflexed apices.

Length ranges 21 – 64μm and width ranges 12 – 18μm. Striae are uniseriate,

distinctly punctate, and are linear at the valve center, becoming slightly radiate at

the apices; 8 – 9 striae per 10μm. Axial area is of uniform width, with a slightly

ovular swelling near the central raphe endings. Raphe is filamentous.

Remarks: Vanlandingham (1964) lists this species as being significantly smaller than the

illustrations presented. Specimens observed here conform to the larger

micrographs he gives, and the maximum size range of this species is expanded for

diatomite in Grant Co., and may be larger for some rare specimens. 108

Distribution: This species is the most abundant Cymbella found in the Quincy Diatomite.

It is most abundant in the Bottom Crude unit of the Frenchman Hills. It is very

rarely observed in the Gorge Amphitheater and ALP, though it was not counted

in point counts in those more northerly locations.

Cymbella ventricosa (Agardh) Agardh 1830

Plate 14 number 6-7

Basionym: Frustulia ventricosa Agardh 1827

Synonyms: Encyonema ventricosum Agardh 1830, Cymbella affinis Kutzing 1844,

Cocconema ventricosum Hassall 1844, Cymbella trugida f. minor Skvortzow

1937.

Type Locality: “Hope fontem Bernardi ad Carlsbad, Bohemia”

Description: Valves are cymbelloid with broadly rounded apices. Ventral side of the

valve is very straight, with the opposite side of the valve being strongly convex.

Length ranges 12 – 21μm and width ranges 5 – 7μm. Striae are uniseriate with

somewhat distinct punctate, and are slightly to strongly radiate through the valve;

11 – 14 striae per 10μm. Axial area is of uniform width and is slightly wider

around the. Raphe is filiform.

Remark: The small size, straight ventral side of the valve, and the comparatively wider

axial area surrounding raphe are the best factors in distinguishing this species

from C. cistula.

Distribution: This species of Cymbella is particularly rare. It is found only in the Bottom

Crude of the Frenchman Hills, being absent in all other units and localities. 109

Genus: Placoneis Mereschkowsky 1903

Placoneis near gastrum (Ehrenberg) Mereschkowsky 1903

Basionym: Pinnularia gastrum Ehrenberg 1843

Synonyms: Schizonema gastrum (Ehrenberg) Kuntz 1898

Description: Valves are elliptical lanceolate to rhombic lanceolate with protracted

terminated apices. Length ranges 21 – 40μm and width ranges 9 – 14μm. Striae

are uniseriate and radiate through the valve. Punctae are not distinct in most

specimens, with larger specimens sometimes showing more clearly defined

punctae. Central striae may be alternately short and long, creating a

subrectangular to nearly ovate central area that may also be slightly larger on one

side of the valve. Central striae for some variants may also approach near even,

however striae counts will be consistent; 9 – 11 striae per 10m. Axial area is

very narrow through the valve face. Raphe is straight becoming hooked at the

apices.

Type Locality: Vera Cruz, Mexico. New Haven, Connecticut, USA. Iceland.

Remarks: Cox (2003) provides measurements that are slightly larger than those provided

by Patrick and Reimer. However, the plates she provides and those she presents

from Hustedt (fig. 7 – 32) are in close agreement with the micrographs presented

by Vanlandingham (1964) (plt. 43 and 44), to include shape of the central area

and the variation in valve outline. Cox also cites fossil deposits in Europe as

known localities, but makes no mention of North American variation in the

species. Fossil variation is presumed to include variation in central striae length

as observed herein. The specimens with consistently straight central striae and a 110

more ovate central area are likely a different species. However, due to the rarity

of these specimens, a complete examination of all forms is necessary before

designation of a new species is done. Vanlandingham (1964) called all similar

species with variable central striae P. gastrum. This does not fit the definition

proposed by Cox (2003), but does fit some of the illustrations of Ehrenberg

(1843) shown by Cox (2003) (figures 4-5, p.58). Due to the variation in central

striae, the use of P. near gastrum is used herein, with further examination of this

species recommended.

Distribution: This is the very rare in the Quincy Diatomite. It is found in the Frenchman

Hills, rarely in the Gorge Amphitheater, and appears to be absent in the ALP

Placoneis rostrata (Mayer) Cox 2003

Plate 11 number 3-5

Basionym: Navicula dicephala var. rostrata Mayer 1917

Synonyms: Navicula elginensis var. rostrata (A. Mayer) Patrick in Patrick & Reimer v.1

1966.

Description: Valves are linear lanceolate to lanceolate with capitate apices, often being

nearly apiculate. Length ranges 25 – 50μm, with width ranging 14 – 25μm.

Striae are distinctly punctate and uniseriate, strongly radiate throughout the valve, and have 7 – 9 striae per 10 μm. Central area has irregularly shortened striae,

creating a subrectangular to oval shaped central area. Raphe is straight and

filamentous, with terminal ends being hooked towards one side of the valve. 111

Proximal ends of the raphe are distinctly enlarged, being approximately double

the punctae size.

Type Locality: Hall. Stavsinge, Holmgärde. England. Fossil.

Remarks: Specimens observed in the Quincy Diatomite had apices more commonly

subcapitate. Central area exhibits varying degrees of angularity, with this

variation being independent of the size of the specimen. Quincy Diatomite

species are also smaller than the measurements provided by Cox (2003), and the

central area may be less distinctly rectangular than Cox (2003) describes. Striae

counts and patterns otherwise match, Thomas 1981 designated specimens as

Navicula amphibola (Figure 12, 6, p. 42). These figures match the designation

for P. rostrata, and are thus considered a synonym here, as this paper was used

for identification purposes.

Distribution: This species is seen rarely across all sampling localities, and is equally

abundant in all three units.

Placoneis elginensis (Gregory) Cox 2003

Plate 11 number 6

Basionym: Pinnularia elginensis Gregory 1856

Synonyms: Navicula elginensis (Gregory) Ralfs in Pritchard 1861

Description: Valves are linear lanceolate with subcapitate apices. Length ranges 30 –

32μm and width ranges 8 – 10μm. Striae are distinctly punctate, radiate

throughout the valve, and uniseriate; 11 – 12 striae per 10μm. Central area is

created by shortened striae forming a rectangular to bow-tie shaped area. Raphe 112

is straight, with a narrow axial area. Terminal fissures of raphe are hooked

towards one side of the valve.

Remarks: Cox (2003) discusses this species based on central area and valve outline, but

neglects to give it a complete range of length as she breaks it out from its allies.

It’s occurrence in this material is smaller in length than provided, and thus the

measurements provided include the maximum listed by Cox and the minimum

observed.

Cox (2003) also figures Placoneis sp. (fig. 68, 69). This species matches the

lower end measurements and match the nominal striae count of P. elginensis.

These species are thus considered part of P. elginensis in this material and

account for the slightly smaller length and breadth measurements provided.

Distribution: This species is very rare, having been observed only twice in the Frenchman

Hills in the Bottom Crude unit. It was not observed at any other sampling

locality.

Family: Gomphonemataceae Kützing 1844

Genus: Gomphonema Ehrenberg 1832

Gomphonema angustatum

Description: Valves are rather linear, with maximum width being slightly eccentric of the

valve face. Length ranges 39 – 40μ, and width is approximately 6μm. Striae are

straight, becoming slightly curved radiate near the apices of the valve. Striae are

also distinctly punctate, with an isolated punctum at the end of the median striae 113

on one side of the valve; approximately 10 striae per 10μm. Raphe is straight

with the axial area becoming narrower towards the apices.

Remarks: This species was listed by Vanlandingham (1964), but was not assigned to a

particular variant. It is closest morphologically to G. angustatum v. obtusatum

(Kützing) Grunow, however it is longer, and is not attenuated at the apices.

Distribution: This species is very rare; it is only found in the lower portions of the

Bottom Crude unit in the Frenchman Hills.

Gomphonema cholnokyites Vanlandingham 1964

Plate 5 number 4-10

Basionym: Gomphonema cholnokyites Vanlandingham 1964

Synonyms: none

Description: Valves are lanceolate to elliptical lanceolate, with slightly protracted apices.

Length ranges 21 – 25μm and width ranges 6 – 7μm. Striae are straight and

distinctly punctate, with one of the median striae being of reduced length

compared to the rest; 7 – 9 striae per 10μm, with striae density being greatest near

the apices. Raphe is straight with a narrow axial area.

Type Locality: Grant County, Washington, USA. Miocene.

Remarks: Valve outline and initial appearance is similar to G. lanceolatum v. insignis,

but the species is distinguished by its shorter length and the appearance of a

narrower mid-region of the valve, lower striae counts, and a slightly narrower area

around the raphe. 114

Distribution: This species is rare through the Frenchman Hills and Gorge Amphitheater,

but is somewhat common in the ALP.

Gomphonema gracile Ehrenberg emend. Van Heurck 1885

Plates 5 number 1, Plate 6 number 5-9

Basionym: Gomphonema gracile Ehrenberg 1838

Synonyms: Gomphonema gracile var. dichotomum (Kutzing) Van Heurck 1885,

Gomphonema gracile var. naviculoides (Smith) Grunow in Van Heurck 1880,

Gomphonema gracile var. dichotoma Vanlandingham 1964, Gomphonema

lanceolatum Ehrenberg 1841, Gomphonema grunowii Patrick 1975

Description: Valves are lanceolate to linear lanceolate with acutely rounded apices. Pore

field at the end of one apex is occasionally visible in exceptionally well preserved

specimens. Length ranges 28 – 70μm, with width ranging 7 – 13μm. Striae are

gently radiate and distinctly punctate. Medial striation on one side of the valve

has a separate and distinct punctum very close to the long striae. The opposite

striae to the long median is significantly shorter; 8 – 10 striae in 10μm. Axial

area is narrow, becoming slightly broader in larger specimens. Raphe is

filamentous, with terminal ends hooked away from the isolated punctam towards

the ventral side of the valve.

Type Locality: Bei Berlin, vielleicht auch bei Tennstädt in Thüringen. Germany.

Remarks: Counts for this species were done on for the varieties Gomphonema gracile

var. naviculoides (Smith) Grunow and Gomphonema gracile var. dichotomum

(Kutzing) Van Heurck, It was subsequently noted that Patrick and Reimer v. 2 115

(1975) combined these varieties into the single species G. gracile. Thus this

description accounts for both varieties.

G. grunowii is treated herein as a junior synonym to G. gracile; Lange-Bertalot

viewed Ehrenberg’s description of G.gracile (1838) and G. lanceolatum (1841) as

two independent descriptions of the same species. The authors agree with this

opinion, and therefore use the G. gracile name for all species fitting the

descriptions of G. grunowii and G. gracile.

Distribution: This species is very rare. It is found in all sampling localities, but most

common in the Top Crude unit.

Gomphonema affine var. insigne (Gregory) Andrews 1970

Basionym: Gomphonema insignie Gregory 1856

Synonyms: Gomphonema insignie f. minor Grunow in Van Heurck 1880, Gomphonema

lanceolatum var. insignis (Gregory) Cleve 1894.

Description: Valves are lanceolate with protracted tapering to the rounded apices. Length

ranges 25 – 60μm and width ranges 6 – 9μm. Striae are straight and punctate,

with some larger specimens showing radiate striae towards the apices. Median striae has a separate and distinct punctum at the end, with the opposite striae being

shortened to create a clear area in the central portion of the valve face; 8 – 11

striae per 10μm. Raphe is straight, with a narrow axial area that becomes broader

in the valve face center.

Type Locality: Duddingston Loch, Scotland. 116

Remarks: Similar to Gomphonema cholnokyites, but is generally longer, with higher

striae counts and a more broad axial area. Confusion between the two is usually

limited to occurrence of 8 striae per 10μ and approximately 25μ length, in which

case the valve outline become the best diagnostic tool, with G. lanceolatum v.

insignis having a more lanceolate appearance.

Patrick and Reimer (1975) gave a slightly larger size range (30 – 60μm) than

Vanlandingham (1964) (25 – 55μm). The measurements provided herein are a

combination of the measurements given by Vanlandingham and Patrick and

Reimer.

Distribution: This species is rare but evenly distributed across the Frenchman Hills and

Gorge Amphitheater; however it is much more abundant in the ALP.

Gomphonema dichotomum Kützing 1833

Basionym: Gomphonema dichotomum Kützing 1833

Synonyms: Gomphonema intricatum var. dichotoma (Kützing) Grunow 1880,

Gomphonema intricatum var. pumila Grunow 1880.

Description: Valves are linear with a protracted apex at one end and broadly rounded

apex at the other. Length ranges 21 – 25μm, with width ranging 6 – 7μm. Striae are slightly radiate and distinctly punctate. Median stria has a separate and distinct

punctum at the end, with the opposite striae being shortened to create a clear

hyaline area in the central portion of the valve face. Shortened striae may be so

short as to appear completely absent; 12 – 13 striae per 10μm. Axial area is

broad, and is near 1/3 of the valve center. Raphe is straight. 117

Type Locality: “An Conferva glomerata und Conferva globulin aim Bruchteiche bei

Ternstädt in Thüringen…”, Germany.

Remarks: Length given here is smaller than that given by Patrick and Reimer v. 2 (1975),

however these measurements match those given by Vanlandingham (1964) and

illustrations from both authors.

Distribution: This species is very rare across all sampling localities, with very few

specimens encountered.

Genus: Gomphosphenia Lange-Bertalot 1995

Gomphosphenia grovei (Schmidt) Lange-Bertalot 1995

Plate 5 number 2-3

Basionym: Gomphonema grovei Schmidt 1899

Synonyms: none

Description: Valves clavate in outline, with a broadly rounded apex and a subrostrate

apex. Length ranges 15 – 25μm, with width ranging 6 – 9μm. Striae are

distinctly punctate, and radiate through most of the valve face, becoming straight

near the subrostrate apex. Striae are irregularly long, with discontinuous ends

near the valve face center creating a largely hyaline valve face. Discontinuous

striae ends may create a second, inner row of punctae in the central portion of the

valve face; 9 – 12 striae per 10μm. Raphe is straight.

Type Locality: Pit River, Shasta County (comm. Grove), California, Washington County,

fossil, USA. 118

Remarks: This species is among the most distinctive in the Quincy Diatomite flora. Its

large hyaline central area is most diagnostic. Size range for the fossil species is

slightly smaller than those given by Patrick and Reimer v. 2 (1975) for modern

specimens.

Distribution: Seen rarely across all sampling localities, though it is most common in the

Frenchman Hills in the Four Crude unit.

Order: Achnanthales Silva in Lewin 1962

Family: Achnanthaceae Kützing 1844

Genus: Planothidium Round and Bukhtiyarova 1996

Planothidium conspicuum Aboal 2003

Plate 7 number 1-6

Basionym: Achnanthes conspicua Mayer 1919

Synonyms: none

Type Locality: Bayern, near Reichenhall, Germany.

Description: Valve outline is elliptical, with some specimens showing a subrectangular

outline due to straighter valve sides. Length ranges 7 – 10μm and width ranges 4

– 5μm. Striae are linear at the mid portions of the valve, and become radiate

towards the apices, with 12– 14 striae per 10μm. Punctae are indistinct under

LM, and appear doubly punctate under SEM. Araphid valve has a broad,

lanceolate pseudoraphe. One side of the valve also shows a gap in the striae,

corresponding to the raphid valve. Raphid valve has a very narrow axial area

around the raphe. The gap in striae on the P valve is corresponds directly with 119

that on the R valve. However this gap oscillates between readily discernable to

highly obscure on the raphid valve.

Distribution: This species is rare in the Frenchman Hills and Gorge Amphitheater, and is

absent in the ALP. It is most common in the middle of the Bottom Crude unit. It

is the most common species of Planothidium in the Quincy Diatomite.

Remarks: This species is most easily distinguished from P. ellipticum based on the large

pseudoraphe, the consistently smaller size, and the absence of the “U” shaped,

hooded clear area.

Planothidium ellipticum (Cleve) Edlund 2001

Plate 7 number 10 &14-15, Plate 21 number 2

Basionym: Achnanthes lanceolata var elliptica Cleve 1891

Synonyms: Achnanthidium lanceolatum var. ellipticum (Cleve) Meister 1912,

Planothidium ellipticum (Cleve) Round and Bukhtiyarova 1996

Type Locality: Åbo, Finland.

Description: Valve outline is elliptical to nearly circular in smaller specimens.

Length ranges 10 – 15μm and width has a range of 6 – 7μm. Striae are strongly

radiate at the apices, becoming gentler in the mid-valve region. Striae punctae are

often obscure in LM, and appear rectangular under SEM. Valves have 13-14

striae per 10μm. Araphid valve has a narrow pseudoraphe that is linear and raised

on the interior of the valve. A “U” shaped clear area is present on one side of the

P valve side and is hooded. The hood partially covers the clear area opening,

creating a small undulation in LM. The raphid valve has a narrow axial area 120

around the raphe. A gap in striae exists on one side of the R valve corresponding

to the clear area on the P valve.

Distribution: This species is rare in the Frenchman Hills and Gorge Amphitheater, and is

absent in the ALP. It is most common in the middle of the Bottom Crude unit.

Remarks: This species is most easily distinguished from P. conspicuum based on the

smaller, narrower pseudoraphe, the consistently larger size, and the presence of

the “U” shaped, hooded clear area.

Planothidium lanceolatum (Brébisson ex Kützing) Lange-Bertalot 1999

Plate 21 number 1

Basionym: Achanthidium lanceolatum Brébisson ex Kützing 1846

Synonyms: Achnanthes lanceolata (Brébisson ex Kützing) Grunow 1880,

Achnantheiopsis lanceolata (Brébisson ex Kützing) Lange-Bertalot 1997

Description: Specimens have a lanceolate to sub-elliptical valve outline. Length ranges

12 – 13μm, width 6 – 7μm. Striae are linear in the central valve area, becoming

gently radiate towards the apices. Striae are robust and rectangular shaped

doubly punctate rows, with punctae sometimes being poorly divided, and

appearing singular. Punctae are indistinct under LM. Araphid valve has a

narrow, lanceolate shaped pseudoraphe that becomes broader in the central area.

There is also a horseshoe shaped, hooded clear area on one side of the valve. The

hood is strongly silicified and is visible in LM. Raphid valve was not observed in

light microscopy. R valve shows a narrow raphe slit with a narrow axial area.

Type Locality: Calvados, Falaise, France 121

Remarks: Lanceolate shape is the most distinguishing feature of this diatom, along with

punctae shape.

Distribution: This species was only observed once in LM and once in SEM, and is the

rarest of all the Planothidium species in this deposit.

Planothidium rostratum (Østrup) Lange-Bertalot 1999

Plate 7 number 7-9 & number 11-12

Basionym: Achnanthes rostrate Østrup 1903

Synonyms: Achnanthes lanceolata var. rostrata Hustedt 1911, Achnanthes lanceolata

ssp. rostrata (Østrup) Lange-Bertalot 1993.

Description: Valves have a rhombic lanceolate outline with protracted rostrate apices.

Length ranges 15 – 32μm, and width ranges 6 – 10μm. Striae are radiate,

becoming strongly so near the apices. Striae are also very robust, singly punctate,

and have 11 – 13 striae per 10μm. Punctae are rectangular, and may be seen in

LM. Araphid valve has a narrow pseudoraphe, with a horseshoe shaped and

hooded clear area on one side of the valve. Raphid valve has a narrow axial area

around the raphe, with a series of irregularly shortened striae on the side of the P

valve clear area. Central area is large and hyaline.

Type Locality: Koh Chang: Klong Sarlakpet, Thailand.

Remarks: Specimens assigned to this species are significantly larger than the

measurements given by Vanlandingham (1967), who restricts the length to 23μm.

Based on new samples taken from the original sampling sites illustrated, it is 122

believed these specimens were deeper in the section than Vanlandingham had

access to.

Distribution: This species is most abundant in the Frenchman Hills, and is absent from

the Gorge Amphitheater and ALP. It is seen most commonly in the Bottom

Crude in the same level in the Bottom Crude where the other Planothidium

species are most abundant.

Order: Bacillariales Hendey 1937

Family: Bacillariaceae Ehrenberg 1831

Genus: Nitzchia Hassall 1845

Nitzschia amphibia Grunow 1862

Plate 13 number 8

Basionym: Nitzschia amphibia Grunow 1862

Synonyms: none

Description: Valves are linear lanceolate to rhombic lanceolate with protracted rostrate

apices in the larger specimens. Length ranges 12 – 25μm, with width ranging 3 –

4μm. Striae are linear, extending across the valve face to the raphe. Punctae on

striae are indistinct in LM with 15 – 17 striae per 10μm. Raphe follows the valve

margin, is continuous from apex to apex, and is distinguished by subtending

fibulae consistently sized 1 – 2μm apart. Marginal ridges are also visible in

LM.

Type Locality: Austria. 123

Remarks: This is the only species of Nitzschia observed in the Quincy Diatomite. Most

valves observed were lightly silicified.

Distribution: This species is restricted to the Frenchman Hills, being found only in the

Bottom Crude and Four Crude units. 124

Extremely Rare Species List:

The remaining species listed were observed and identified. All species listed here are extremely rare (less than 5 observations over all samples; most less than 2), with many not counted in point counts and few in position for light micrograph.

Fragilaria capucina var. lanceolata Grunow 1881

Staurosira construens var. binodis Cleve 1882. Plate 15 number 6

Eunotia clevei Grunow 1891. Plate 21 number 7

Eunotia pseudopectinalis Hustedt 1924

Cocconeis sp. Plate 21 number 8

Achnanthidium exiguum (Grunow) Czarnecki 1994

Navicula avenacea (Rabenhorst) Brébisson ex Grunow 1878

Navicula near insula Metzelin and Lange-Bertalot 1998

Navicula pantocsekiana DeToni1891

Navicula seminuloides Hustedt 1937

Craticula sp. Grunow 1868

Sellaphora sp. aff. americana (Ehrenberg) Mann 1989. Plate 21 number 10

Sellaphora laevissima (Kützing) Mann 1989. Plate 21 number 9

Stauroneis phoenicenteron (Nitzsch) Ehrenberg 1845. Plate 13 number 3

Stauroneis near acuta Smith 1853. Plate 13 number 4

Rhoicosphenia curvata (Kützing) Grunow ex. Rabenhorst 1864

Placoneis amphibola (Cleve) Cox 2003

Placoneis anglica (Ralphs) Cox 2003. Plate 11 number 7 125

Placoneis placentula Ehrenberg 1843. Plate 11 number 1-2

Gomphonema acuminatum var. montanum (Schumann) Grunow in Van Heurck 1880.

Plate 6 number 10-11

Gomphonema tenellum Kützing 1844. Plate 6 number 1-4

Reimeria sinuata (Gregory) Kociolek and Stoermer 1987. Plate 12 number 12

126

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140

III. Plate Captions

Plate 1: 1 – 9. Frenchman Hills, core Q0412-11. A. granulata. 1 – 9. 1. depth 7.0m. 2.

depth 4.0m. 3. depth 9.4m. 4. Diagenetically modified A. granulata chain.

Depth 6.2m. 5. depth 8.2m. 6. Resting frustule, depth 6.9m. 7. Diagenetically

modified frustule, depth 4.0m. 8. Chain of frustules, depth 9.4m. 9. depth 4.0m.

10 – 12. A. agassizi. 10. Auxospore chain, focus on outer frustule, depth 8.8m.

11. depth 6.2m. 12. Auxospore chain, focus on inner frustule, depth 8.8m. Scale

bars 20μm. 141 142

Plate 2: 1 – 10. A. canadensis. Frenchman Hills, core Q0412-11. 1. depth 37.3 ft. 2.

depth 6.2m. 3. depth 8.8m. 4. depth 9.4m. 5. depth 7.5m. 6-7. depth 10.4m. 8.

depth 4.0m. 9. depth 4.0m. 10. depth 10.4m . Scale bars 20μm except 6, 7, 9, 10,

with 10μm scale bars. 143 144

Plate 3: 1 – 5. Ellerbeckia baileyi. 1-4. Frenchman Hills, core Q0412-11, depth 5.7m. 5.

Frenchman Hills, core Q0412-11, depth 5.5m. Scale bars 20μm. 145 146

Plate 4: Melosira undulata. 1a. Outer part of frustule. Frenchman Hills, core Q0412-11,

depth 8.8m. 1b. Focus on inner frustule. Frenchman Hills, core Q0412-11, depth

5.5m. 2. Frenchman Hills, core Q0412-11, depth 5.5m. Scale bar 10μm. 147 148

Plate 5: Gomphonema gracile. 1. Ancient Lakes Park, depth 1.0m. 2 – 3.

Gomphosphenia grovei. Frenchman Hills, core Q0412-11, depth 5.5m. 4 – 10

Gomphonema cholnokytes. 4. Frenchman Hills, core Q0412-11, depth 5.5m. 5.

Ancient Lakes Park, depth 1.0m. 6. Ancient Lakes Park, depth 2.5m. 7-8.

Ancient Lakes Park, depth 1.0m. 9. Ancient Lakes Park, depth 2.0m. 10. Ancient

Lakes Park, depth 1.0m. Scale bar 10μm. 149 150

Plate 6: G. tenelum. 1 – 4. 1 & 2. Ancient Lakes Park, depth 2.5m. 3. Ancient Lakes

Park, depth 2.0m. 4. Frenchman Hills, core Q0412-11, depth 5.5m. G.

gracile. 5 – 9. 5, 6, &7. Ancient Lakes Park, depth 2.5m. 8. Frenchman Hills,

core Q0412-11, depth 5.5m. 9. Ancient Lakes Park, depth 1.0m. G.

accuminatum var. montanum. 10 & 11 Ancient Lakes Park, depth 1.0m. Scale

bars 20μm, except 2 and 3, with 10μm scale bars. 151 152

Plate 7: P. conspicuum. 1 – 6. 1. Araphid valve. Frenchman Hills, core Q0412-11,

depth 5.5m. 2. Raphid valve. Frenchman Hills, core Q0412-11, depth 35.2 ft.

3. Araphid valve. Frenchman Hills, core Q0412-11, depth 5.5m. 4. Raphid valve.

Frenchman Hills, core Q0412-11, depth 5.5m. 5-6. Araphid valve. Frenchman

Hills, core Q0412-11, depth 5.5m. P. rostratum. 7 – 9 and 11 – 12. 7 – 9

Bottom Crude, Frenchman Hills. 7. Araphid valve. Frenchman Hills, core

Q0412-11, depth 5.5m. 8. Raphid valve. Frenchman Hills, core Q0412-11, depth

5.5m. 9. Araphid valve. Frenchman Hills, core Q0412-11, depth 5.5m. 11-12.

Araphid valve. Frenchman Hills, core Q0412-11, depth 5.5m. P. ellipticum. 10 a

& b and 14 – 15. 10. Frenchman Hills, core Q0412-11, depth 5.5m. a. Araphid

valve. b. Raphid valve. 13. Araphid valve. Frenchman Hills, core Q0412-11,

depth 5.5m. 14. Raphid valve. Frenchman Hills, core Q0412-11, depth 5.5m.

Scale bars 10μm, except 4, 5, have 5μm scale bars. 153 154

Plate 8: Tetracyclus williamsensii. 1-2. Frenchman Hills, core Q0412-11, depth 5.5m.

3. Frenchman Hills, core Q0412-11, depth 4.5m. 4. Frenchman Hills, core

Q0412-11, depth 5.5m. 5. Frenchman Hills, core Q0412-11, depth 4.5m. 6.

Frenchman Hills, core Q0412-11, depth 5.5m. Scale bars 10μm. 155 156

Plate 9: 1. T. ellipticus. Frenchman Hills, core Q0412-11, depth 5.5m. 2. T. linearis.

Ancient Lakes Park, depth 2.5m. 3. T. lacustris. Frenchman Hills, core Q0412-

11, depth 6.2m. Scale bars 10μm. 157 158

Plate 10: 1 – 2. Tetracyclus polygibbum Frenchman Hills, core Q0412-11, depth 4.5m. 3.

T. lata. Frenchman Hills, core Q0412-11, depth 2.1m. 4. Pinnularia nodosa.

Ancient Lakes Park, depth 0.0m. 5. Sellaphora bacillum. Frenchman Hills, core

Q0412-11, depth 5.5m. 6. Meridion circulare. Ancient Lakes Park, depth 2.5m.

Scale bars 10μm. 159 160

Plate 11: Placoneis placentula 1 – 2. 1. 2. Frenchman Hills, core Q0412-11, depth 1.5m.

3 – 5 P. rostrata. 3- 4. Frenchman Hills, core Q0412-11, depth 5.5m. 5.

Frenchman Hills, core Q0412-11, depth 4.0m. 6. P. elginensis. Frenchman Hills,

core Q0412-11, depth 4.5m. 7. P. anglica. Frenchman Hills, core Q0412-11,

depth 5.5. 8. Diatoma anceps. Frenchman Hills, core Q0412-11, depth 4.5m. 9.

Eunotia curvata. Ancient Lakes Park, depth 2.5m. Scale bars 10μm. 161 162

Plate 12: 1 – 5 Eunotia veneris. 1. Frenchman Hills, core Q0412- 11, depth 5.5m. 2.

Frenchman Hills, core Q0412-11, depth 5.6m. 3-4. Ancient Lakes Park, depth

1.0m. 5. Frenchman Hills, core Q0412-11, depth 4.5m. 6. E. pectinalis. Ancient

Lakes Park, depth 2.5m. 7. E. pectinalis var. minor. Ancient Lakes Park, depth

2.5m. 8 – 11 Diatoma hyemale var. mesodon. 8. Ancient Lakes Park, depth

2.5m. 9. Frenchman Hills, core Q0412-11, depth 35.2 ft. 10. Ancient Lakes Park,

depth 2.5m. 11. Ancient Lakes Park, depth 2.5m. 12. Reimeria sinuata.

Frenchman Hills, core Q0412-11, depth 4.5m. Scale bars 10μm. 163 164

Plate 13: 1 – 2 Pinnularia near major. 1. Ancient Lakes Park, depth 2.0m. 2. Ancient

Lakes Park, depth 0.0m. 3. Stauroneis phoenicenteron. Ancient Lakes Park,

depth 0.0m. 4. Stauroneis near acuta. Ancient Lakes Park, depth 2.0m. 5 – 6

Navicula reimertes. Frenchman Hills, core Q0412-11, depth 5.5m. 7. Navicula

scutelloides. Frenchman Hills, core Q0412-11, depth 4.5m. 8. Nitzchia

amphibia. Frenchman Hills, core Q0412-11, depth 5.5m. Scale bars 10μm,

except 1, 2 have 25μm scale bars. 165 166

Plate 14: 1 – 4 and 10. Cymbella cistula. 1. Frenchman Hills, core Q0412-11, depth

5.5m. 2. Frenchman Hills, core Q0412-11, depth 5.6m. 3. Frenchman Hills, core

Q0412-11, depth 5.5m. 4. Ancient Lakes Park, depth 2.0m. 10. Ancient Lakes

Park, depth 0.0m. 5 and 9. Diploneis ovalis. 5. Frenchman Hills, core Q0412-11,

depth 5.5m. 9. Frenchman Hills, core Q0412-11, depth 5.5m. 6 – 7 Cymbella

ventricosa. Frenchman Hills, core Q0412-11, depth 5.5m. 8. Cavinula

pseudoscutiformis. Frenchman Hills, core Q0412-11, depth 5.6m. Scale bars

10μm. 167 168

Plate 15: 1. Pseudostaurosira brevistriata. Frenchman Hills, core Q0412-11, depth

5.5m. 2 – 3 P. brevistriata var. subcapitata. 2. Frenchman Hills, core Q0412-11,

depth 5.6m. 3. Frenchman Hills, core Q0412-11, depth 5.5m. 4 – 5 Staurosira

construens. Frenchman Hills, core Q0412-11, depth 5.5m. 6. S. construens var.

binodis. Frenchman Hills, core Q0412-11, depth 5.5m. 7 & 10 Fragilaria

lapponica. 7. Frenchman Hills, core Q0412-11, depth 5.5m. 10. Frenchman

Hills, core Q0412-11, depth 5.5m. 8. Staurosirella pinnata var. lancettula.

Frenchman Hills, core Q0412-11, depth 5.5m. 9 and 14. Fragilaria near socia.

9. Frenchman Hills, core Q0412-11, depth 5.5m. 11 – 12 Fragilaria near

lenoblei. 11. Frenchman Hills, core Q0412-11, depth 35.2 ft. 12. Frenchman

Hills, core Q0412-11, depth 5.5m. 13 and 15 – 19. Fragilariforma virescens. 13.

Ancient Lakes Park, depth 1.0m. 15. Ancient Lakes Park, depth 1.0m. 16 –18.

Ancient Lakes Park, depth 1.0m. 19. Ancient Lakes Park, depth 2.5m. Scale bars

10μm. 169 170

Plate 16: 1 – 16. Staurosira construens var. venter. Size gradational series. All

specimens are from the Frenchman Hills, core Q0412-11, depth 5.5m. Scale bars

10μm, except 1 – 5, 7, 8, with 5μm scale bars. 171 172

Plate 17: 1 – 5 Actinocyclus motilis. Size gradational series. 1. Frenchman Hills, core

Q0412-11, depth 3.8m. a. Focus on upper valve. b. Focus on inner valve

showing labiate processes. 2. Frenchman Hills, Frenchman Hills, core Q0412-11,

depth 3.9m. 3-5. Frenchman Hills, core Q0412-11, depth 3.9m. Scale bars 10μm. 173 174

Plate 18: 1 – 3. Actinocyclus krasskei. 1-2. Frenchman Hills, core Q0412-11, depth

3.8m. 3. Frenchman Hills, core Q0412-11, depth 3.9. 4. Actinocyclus sp. SEM

showing upper valve surface and areola. Ancient Lakes Park, depth 1.5m. 5.-6.

A. motilis. 5. SEM showing inner valve area. Ancient Lakes Park, depth 1.0m.

6. SEM showing upper valve surface. Ancient Lakes Park, depth 2.0m. Scale

bars 10μm. 175 176

Plate 19: 1 – 7. Resting cells from Ancient Lakes Park, depth 1.0m. 1-2, 4-5.

Actinocyclus (?) resting spores. 1a. Outer valve wall. 1b. Inner valve. 2a. Outer

valve wall. 2b. Inner valve wall. 4a. Outer valve wall. 4b. Inner valve wall. 4c.

Focus through specimen showing encircling ring of silica on valve face. 5. Chain

of resting stage cells interior view. 3 and 7. Meridion circular internal valves.

3a. Outer valve wall. 3b. Inner valve bordered by outer valve. 3c. Internal valve

interior. 7a. Outer valve wall in girdle view. 7b. Focus on outer and inner valves.

7c. Internal valve within epivalve. 6. Aulacoseira canadensis auxospore chain. a.

Outer valve wall, showing pore shape. b. Interior of valve. 177 178

Plate 20: 1. Tetracyclus lancea. Frenchman Hills, core Q0412-11, depth 5.5m. a. Upper

valve surface. b. Inner valve. 2-5. T. williamsensii. 2. Girdle band with

characteristic “ox-bow” shape. Frenchman Hills, core Q0412-11, depth 5.5m. a.

Upper view of septum. b. Lower focus on septum. 3. SEM, inner valve showing

costae structure and apical pore-field. Frenchman Hills, core Q0412-11, depth

9.4m. 4. SEM showing single rimoportulae on valve face near mantle.

Frenchman Hills, core Q0412-11, depth 10.2m. 5. SEM showing apical pore-field

and rimoportulae. Frenchman Hills, core Q0412-11, depth 5.5m. 6 – 7 Melosira

undulata. Frenchman Hills, core Q0412-11, depth 6.9m. 6. SEM showing

undulatory inner valve structure and through-going pores. 7. SEM showing outer

vale wall structure and ringleist. Scale bars 10μm. 179 180

Plate 21: 1. Planothidium lanceolatum. Biseriate punctae with poor silicification.

Frenchman Hills, core Q0412-11, depth 8.4m. 2. Planothidium ellipticum.

Araphid valve. Frenchman Hills, core Q0412-11, depth 4.1m. 3. Neidium

near iridis var. ampliatum. Ancient Lakes Park, depth 2.5m. 4 & 5. E. baileyi.

Frenchman Hills, core Q0412-11, 4. depth 8.4m 5. depth 4.4m. 6.

Pseudostaurosira brevistriata var. subcapitata. Frenchman Hills, core Q0412-11,

depth 6.1m. 7. Eunotia clevei. Frenchman Hills, core Q0412-11, depth 4.0m. 8.

Cocconeis sp. Frenchman Hills, core Q0412-11, depth 5.5m. a. Raphid valve,

upper focus. b. Araphid valve, lower focus. 9. Sellaphora laevissima.

Frenchman Hills, core Q0412-11, depth 4.0m. 10. Sellaphora near americana.

Frenchman Hills, core Q0412-11, depth 9.4m. 11–12. Fragilariforma

vaucheriae. Frenchman Hills, core Q0412-11, depth 5.5m. Scale bars 10μm. 181 182

Plate 22: Fragilariforma intortus. Focus series through three specimens. Frenchman

Hills, core Q0412-11, depth 7.5m. 1-3. small length specimen showing break in

striae and spines. 4-5. showing large apical pore field. 6-10. focus looking

through interior of valve to exterior, with striae break being most distinct in the

central striae. Scale bars 10μm. 183 184

Plate 23: 1-8. Fragilariforma intortus. Size gradational series. Frenchman Hills, Core

Q0412-11. 1 & 3-4. depth 6.9m. 2 and 5. depth 5.5m. 6. depth 9.4m. 7. depth

5.6m. 8. depth 5.5m. 185 186

Appendix 1 – Sample Numbers and Sampled Cores

List of sample numbers and where the samples were taken from. Samples used on correspondence analysis are also given number designations. 187

SampleNumbers Corenumber Core/SampleLocation 07AM0501 N/A FrenchmanHills 07AM0502 N/A FrenchmanHills 07AM0503 N/A FrenchmanHills 07AM0504 N/A FrenchmanHills 07AM0505 N/A FrenchmanHills 07AM0506 N/A FrenchmanHills 07AM0507 N/A FrenchmanHills 07AM0508 N/A FrenchmanHills 07AM0509 N/A FrenchmanHills 07AM0510 N/A FrenchmanHills 07AM0511 N/A FrenchmanHills 07AM0512 N/A FrenchmanHills 07AM0513 N/A FrenchmanHills 07AM0514 N/A FrenchmanHills 07AM0515 N/A FrenchmanHills 07AM0516 N/A FrenchmanHills 07AM0517 Q041211 WorldMinerals 07AM0518 Q041211 WorldMinerals 07AM0519 Q041211 WorldMinerals 07AM0520 Q041211 WorldMinerals 07AM0521 Q041211 WorldMinerals 07AM0522 Q041211 WorldMinerals 07AM0523 Q041211 WorldMinerals 07AM0524 Q041211 WorldMinerals 07AM0525 Q041211 WorldMinerals 07AM0526 Q041211 WorldMinerals 07AM0527 Q041211 WorldMinerals 07AM0528 Q041211 WorldMinerals 07AM0529 Q041211 WorldMinerals 07AM0530 Q041211 WorldMinerals 07AM0531 Q041211 WorldMinerals 07AM0532 Q041211 WorldMinerals 07AM0533 Q041211 WorldMinerals 07AM0534 Q041211 WorldMinerals 07AM0535 Q041211 WorldMinerals 07AM0536A Q041211 WorldMinerals 07AM0536B Q041211 WorldMinerals 07AM0537 Q041211 WorldMinerals 07AM0538 Q041211 WorldMinerals 07AM0539 Q041211 WorldMinerals 188

07AM0540 Q01209 WorldMinerals 07AM0541 Q01209 WorldMinerals 07AM0542 Q01209 WorldMinerals 07AM0543 Q01209 WorldMinerals 07AM0544 Q01209 WorldMinerals 07AM0545 Q01209 WorldMinerals 07AM0546 A012015 WorldMinerals 07AM0547 N/A GorgeAmphitheater 07AM0548 Q0424 WorldMinerals 07AM0549 Q0424 WorldMinerals 07AM0550 Q0424 WorldMinerals 07AM0551 Q0424 WorldMinerals 07AM0552 Q0424 WorldMinerals 07AM0553 Q0424 WorldMinerals 07AM0554 Q0424 WorldMinerals 07AM0555 Q047161 WorldMinerals 07AM0556 Q047161 WorldMinerals 07AM0557 Q047161 WorldMinerals 07AM0558 Q047161 WorldMinerals 07AM0559 Q047161 WorldMinerals 07AM0560 Q047161 WorldMinerals 07AM0561 Q047161 WorldMinerals 07AM0562 Q047161 WorldMinerals 07AM0563 Q031623 WorldMinerals 07AM0564 Q031623 WorldMinerals 07AM0565 Q0191 WorldMinerals 07AM0701 N/A ALP 07AM0702 N/A ALP 07AM0703 N/A ALP 07AM0704 N/A ALP 07AM0705 N/A ALP 07AM0706 N/A ALP 07AM0707 N/A ALP 07AM0708 N/A ALP 07AM0709 N/A ALP 07AM0801 00BW14 USACE 07AM0802 00BW14 USACE 07AM0803 00BW14 USACE 07AM0804 00BW06 USACE 07AM0805 00BW06 USACE 07AM0806 99BW15 USACE 07AM0807 99BW15 USACE 07AM0808 99BW15 USACE 189

CorrespondenceAnalysisplot# N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 190

N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A 1 2 3 4 5 6 7 8 N/A N/A N/A N/A N/A N/A N/A N/A 191

Appendix 2 – Specimen Picture X-Y Poordinates

X – Y coordinates of species photos from Olympus BX-51 microscope at UNR.

All light micrographs taken, including those not used for plates, are listed. 192

Slides Species position (ft) Stratigraphic Pores Big sp Actinocyclus pores small sp Actinocyclus type mesh Wiery sp Actinocyclus Anomoeoneis sphaerophora agassizi Aulacoseira Aulacoseira canadensis Aulacoseira granulata Aulacoseria granulata diagenetic Craticula sp. cistula Cymbella Q0412-11 07AM05-19a 0.0 12 3004 371883100 07AM05-20a 0.9 0100141280100 07AM05-21a 2.8 01001 0298000 07AM05-22a 4.7 0000210279000 07AM05-23a 6.6 00009122168000 07AM05-24a 8.5 97 2200 68131000 07AM05-25a 8.6 146 5 2 0 11 30 95 3 0 0 07AM05-26a 9.0 20 0 0 1 22 80 112 6 0 0 07AM05-27a 10.9 25 0007 4976004 07AM05-28a 13.4 70000 1253103 07AM05-29a (sandy horizon) 14.0 00000 0 0 000 07AM05-30a (3 row 387 ct) 14.2 70000 0100009 07AM05-31a (++ broken up) 14.7 14 0001 2108002 07AM05-32a 14.9 31301 8 31910 07AM05-33a 16.3 103 0404 21431501 07AM05-34a (++broken up) 18.8 11505 13225101 07AM05-35a 20.6 138 22 0 0 1 8 90 5 0 1 07AM05-36Aa (min. presv) 23.0 44001 2237000 07AM05-36Ba 23.0 166 0300 4104000 07AM05-37a 25.0 62 9002 16195002 07AM05-38a 27.0 37 23 0 0 0 75 144 0 0 1 07AM05-39a 29.8 25 3201118145001 ALP 07AM07-02a 0.0 72001120139300 07AM07-03a 1.6 91000 871571100 07AM07-04a 3.3 20000 38912200 07AM07-05a 4.9 22 0000143104700 07AM07-06a 6.6 18 000016276200 07AM07-07a 8.2 18 020213192400 07AM07-08a 9.9 82002 98185000 11 00000000000000 01010 00100000000000 00260 00100200000000 00330 00000100000100 02010 03500700000400 00100 01000000000000 02300 01020000000001 01100 00000000000100 00010 00100000000200 00110 00000000000100 00010 90000000000000 00000 00000010115193004000 40000000202131000220 00000000001272002000 50000100001110 00000 000002241000010000040 1400000303027121225011 3002013000003203054924 00000000000000 00000 0101017500000020092500101001142001000010300 00000020005264221020 00000000002000 00010 00000000000000 00000 00000000000000 00000 00000000000000 00000 00000000000000 00000 00000000000000 00000 10040000000200000010 Cymbella ventricosa

Diatoma anceps

Diatoma hemalie

Diatoma hiemale v mesodon

Diploneis ovalis

Ellerbeckia balleyi

Eunotia lunaris

Eunotia pectinalis

Eunotia pseudopectinalis

Eunotia pectinalis var minor

Eunotia veneris

Pseudostaurosira brevistriata

Pseudostaurosira brevistriata var elliptica

Pseudostaurosira brevistriata var subcapitata

Fragilaria capucina var laceolata

Staurosira construens

Staurosira construens var venter

Fragilaria lapponica

Fragilaria near lenoblei 193 00000000000100000000 0022000200022001400000020000200005000101000010000200002000410000151300152000192009000000192003000030001010000800030000700001100 00010000000000000000 06000000000000300000 03010000000000200000 01210000000000000001 02100000050100100101 00320000000100010000 42400000010100000100 13310000000020300000 02100000000001000000 30100500000001001014 41110350000411002203 00000000000000000000 21001000000000000001 00010000000000101002 11100000000110200001 00000000100000010002 00000000000000000000 00000000000000100000 00000000000000000000 00000000000000000000 00000000000000000002 00300000000000300001 Fragilaria leptostauron

Fragilaria pinnata var laceolata

Fragilaria virescens

Fragilariopsis (Fragilaria?) spVanlandingham 1964

Gomphonema acuminatum var montanum

Gomphonema angustatum

Gomphonema cholnokytes

Gomphonema gracile v. dichotoma

Gomphonema gracile v naviculoides

Gomphonema grovei

Gomphonema grunowii

Gomphonema affine var insignious

Gomphonema dichotomum

Gomphonema tenellum (?)

Melosira undulata

Meridion circulare

Navicula acceptata

Navicula amphibola

Navicula avenacea

Navicula gastrum 194 0000000 0000000 0000000 0000000 0000000 0004000 0001000 0000000 0000000 0000000 0000000 0000000 0002000 0000000 0000000 0000000 0000000 1010020 0103000 0000000 0000100 0000000 0000000 0000000 0000000 0000000 0000000 0000000 0000000 Navicula near insula

Navicula pantoesekiama

Navicula pseudoscutiformis

Navicula reimertes

Navicula scutelloides

Navicula seminuloides

Neidium iridis

000000000 00000000 00000 00010000 00000 00001000 00000 00000000 00020 00001000 00000 00000000 00000 00001000 00000 00000000 00000 00000000 00000 10000000 00000 00000000 00001 30000100 00000 00000000 00000 00100000 00000 00100000 00000 01010000 32212 00100440 22400901110110 00000 00000000 00100 01000000 00200 01000100 00200 10000002 00000 00000001 00000 00000000 00000 00000000 00000 00000000 00000 00000000 00000 00000000 00000 00000000 Nitzchia amphibia

Opephora glangeaudi

Opephora martyi

Pinnularia major

Pinnularia nodosa

Planothidium conspicuum

Planothidium ellipticum

Planothidium lanceolatum

Planothidium rostratum

Rhoicosphenia curvata

Sellophora bacillum

Stauroneis kriegeri

Synedra tabulata var fasiculata 195 00000011000 00100007000 00100001200 00100101000 00401000000 10000001000 00000000000 00000011000 00001202100 00000211001 00010600110 10000000000 00100010020 00000000000 10010100110 00201411000 21001110021 00000100004 000000000000 01000100010 00013200020 02020200300 00000002000 00000000000 00000000000 00000000000 00000000000 00001000000 00010010000 Synedra near socia (my diagnosis)

Synedra vaucheriae

Synedra sp. (close to fragilariopsis ?)

Tetracyclus ellipticus

Tetracyclus lacustris

Tetracyclus lata

Tetracyclus lancea

Tetracyclus linearis

Tetracyclus polygibbum

Tetracyclus sp H

Unknown sp (with photos, ID post counts) 295/4 250/42 258/31 269/21 171/74 265/29 272/11 294/4 281/16 287/10 286/13 249/39 268/28 251/43 277/17 280/18 139/124 237/81 248/44 272/15 242/54 292/6 300/0 300/0 300/0 300/0 297/0 288/10

Centrics/Araphids ratio (All to ratio of 1 araphid) 196 197

Appendix 3 – Point Count Data

Point count data from counts of the Frenchman Hills and ALP. Counts are listed by species and occurrence within samples. 198

Picturename Slide YCoordinates XCoordinates A.granandcanadensis 0709a 8.4 142.7 Cymbella 0709a 14.3 136.0 Stauroneis 0709a 12.3 140.9 Pinn.Nodosa 0709a 14.4 136.2 M.undulata 0726b 12.8 128.8 Achnantheslanceolata 0526b 13.9 128.8 GomphonemaspA1 0526b 18.1 130.0 NaviculaspB 0526b 18.2 131.3 Tetrasp.affpolygibbum2 0526b 20.2 131.4 Diathiemale2 0526b 20.1 131.1 Naviculabacillum 0526b 20.2 136.1 NaviculoidspB2 0526b 20.2 138.2 NavicspB3 0526b 20.4 139.6 Fragilariasp 0526b 20.5 140.2 Tetrasp.affpolygibbum3 0526b 20.3 140.4 Actin126B 0526b 20.4 142.0 Fragilariasp3 0526b 10.3 134.2 Eunotiaveneris 0526b 10.3 133.4 Largetetra(lata?) 0526b 10.2 129.1 M.undulata2 0526b 14.6 134.9 Eunotia 0526b 14.6 136.9 TetraspH 0527a 14.8 142.1 Tetraspaffpolygibbum4 0527a 14.9 145.5 Tetralata 0527a 13.5 138.6 Eunotialunaris 0527a 10.7 138.6 TetraspH6 0527a 3.3 138.6 TetraDiat2 0527a 5.8 140.3 GomphonemaspB 0527a 4.2 140.9 Gomphonemasp3 0527a 4.1 146.9 GomphonemaspC 0527a 14.6 141.8 Tetraellipticus 0527a 14.3 140.0 CraticulaspA 0527a 10.6 139.3 TetraDiat3 0527a 10.5 138.7 ActinocyclusspE 0528a 3.2 138.8 E.baileyi 0528a 14.4 140.4 araphid5 0528a 13.7 140.4 ActinocyclusspE2 0529b 19.4 147.2 TetraspH2 0530a 18.5 131.9 GomphspD 0530a 18.5 131.9 Tlancea 0530a 18.6 129.0 199

Ebaileyii2 0530a 12.3 130.5 TetraH3 0530a 10.4 130.5 E.baileyi3 0530a 9.3 129.3 Cymbella130A 0530a 5.7 132.1 Naviculabacillum2 0530a 7.3 132.0 NaviculoidB4 0530a 7.3 132.0 Cymbella2 0530a 7.3 130.3 Fragilariasp6 0530a 9.0 132.6 Araphid7 0530a 9.3 132.5 Tetralacustris 0530a 10.2 131.9 T.spH4 0530a 10.2 131.9 Placonthidiumellipticum0530A 0530a 10.1 130.8 GomphonemaC2 0530a 10.4 129.1 TetraDiatheimale 0530a 11.4 128.7 M.undulata 0531b 11.3 132.6 TetraspH5 0531b 11.3 132.6 GomphonemaC3 0531b 11.2 131.1 T.lacust 0531b 11.2 128.8 Araphid8 0531b 11.2 128.8 Araphid9 0531b 11.2 128.6 2Araphid10 0531b 15.4 128.8 Diatomaanceps 0531b 15.5 128.8 NaviculoidspC 0531b 16.9 128.8 E.baileyi4 0531b 21.0 133.4 TetraDiat5 0531b 21.2 133.3 AC010539B 0539b 13.7 144.5 AC020539B 0539b 13.6 144.4 AC030539B 0539b 13.2 143.2 AC040539A 0539a 13.4 142.6 AC050538B 0538b 13.8 141.6 AC060537B 0537b 18.2 137.3 AC070537B 0537b 22.0 140.6 AC080535B 0535b 16.3 138.6 AC090535B 0535b 20.7 137.8 AC100535A 0535a 16.5 139.7 AC110533B 0533b 11.3 143.3 AC120526B 0526b 14.0 134.1 AC130526B 0526b 12.7 130.2 AC140526B 0526b 13.0 144.2 NID010538A 0538a 15.2 141.5 NID020536B2 0536Bb 16.6 134.5 NID040534A 0534a 8.8 142.9 NID050534A 0534a 8.5 143.7 200

NID060532B 0532b 20.0 144.5 NID080531B 0531b 15.6 134.8 NID090530B 0530a 12.8 128.0 NID030538A 0538a 19.0 130.2 NID070532A 0532a 12.8 137.8 NID100531B 0531b 15.7 134.9 NID110531B 0531b 15.9 133.3 NID120531B 0531b 16.1 132.9 NID130527B 0527b 17.2 136.2 NID140527A 0527a 22.4 138.9 NID150527A 0527a 22.4 138.9 NID160527B 0527b 19.0 134.7 NID170526B 0526b 12.9 130.3 MU010537A 0537a 23.1 141.0 MU030537A 0537a 15.2 141.3 MU040536Bb 0536Bb 16.5 131.8 MU050535B 0535b 18.7 140.9 AA010537A 0537a 21.0 143.1 AA050535A 0535a 19.2 139.3 AA080533B 0533b 11.2 141.7 TLac010535A 0535a 16.2 138.1 TLac020533A 0533a 9.4 134.2 TLac030526B 0526b 12.7 130.2 AG010533A 0533a 19.1 132.4 AG020534A 0534a 8.4 142.0 AG030538A 0538A 7.4 130.7 AG040526B 0526b 14.0 134.1 AG050526B 0526b 13.0 128.1 AG060526B 0526b 13.0 129.2 AG070526B 0526b 13.0 134.5 AG080526B 0526b 13.0 137.7 AG090526B 0526b 13.0 140.9 AG100526B 0526b 15.0 146.9 AG110537A 0537a 21.4 143.8 AG120536Aa 0536Aa 12.7 135.7 AG130535B 0535b 18.6 138.2 AG140534B 0534b 21.7 131.2 AG150534B 0534b 21.9 129.6 AG160538A 0538a 14.8 145.8 AG170526B 0526b 12.6 138.1 AG180537A 0537a 21.7 143.9 EB010532B 0532b 20.8 145.8 EB020532B 0532b 21.9 142.0 201

EB030532B 0532b 18.5 143.5 EB040532A 0532a 11.3 137.8 EB050530A 0530a 18.3 134.4 CC010531B 0531b 16.3 141.4 CC020530A 0530a 16.1 136.8 EV010531B 0531b 16.0 133.0 EV020526B 0526b 16.9 135.3 NaviculaspD1 0520a 20.7 141.2 Tetralata2a 0521a 20.4 136.2 Actinocyclus2 0524a 10.7 151.3 Actinocyclus3 0524a 3.2 148.4 TH010527A 0527a 11.2 142.8 TH020527B 0527b 10.8 137.6 NaviculaspE1 0527b 19.4 137.7 Tetralancea1 0530a 14.6 133.7 ActnAl01 0702SEM ActnAl02 0702SEM FragAl01 0702SEM ActnAl03 0704SEM ActnAl04 0704SEM PinnAl01 0704SEM ActnAl05 0707SEM ActnAl06 0707SEM ActnAl07(labiateproc) 0707SEM GphAl07a(wholefrust.) 0707SEM GphAl07b(closeupofpores) 0707SEM ActnAl08(tubeprocess) 0707SEM TlataAl01 0707SEM ActnAl09a(wholefrust.) 0705SEM ActnAl09b(pores) 0705SEM FragAl02 0706SEM TelipAl01 0706SEM TlinAl01 0705SEM CratAl01a(wholefrust) 0705SEM CratAl01b(closeuppores) 0705SEM CratAl01c(proximalraphes) 0705SEM CratAl01d(distalrapheends) 0705SEM TlinAl02 0707SEM TlinAl03 0707SEM NavAl01 0707SEM TetpC101 0536BSEM TlanC101 0537SEM EbalC101 0538SEM 202

MundC101 0534SEM TpolC101 0533SEM EbalC102 0532SEM TspHC101 0531SEM TspHC102 0530SEM TspHC103(withgirdleband) 0530SEM TpsHC104 0530SEM SlopC101 0530SEM AcanC101 0530SEM FragC101 0534SEM MundC102 0534SEM FragC102 0535SEM GompC101 0537SEM TlacC101 0537SEM TlacC102 0537SEM AcanC102 0537SEM TpsHC105 0538SEM TspHC106 0538SEM TspHC107 0538SEM TspHC108 0538SEM TspHC109 0539SEM Danceps 0528a 17.0 138.8 unkacanth1 0530a 13.0 130.5 Ntakei 0530a 13.0 130.2 Npseudoscutiloides 0531a 12.0 141.9 NavAcanth1 0531a 12.0 140.5 Planothidiumconspicuum1a&b 0530a 12.0 141.5 Planothidiumrostratum1a 0530a 12.0 136.9 Placonthidiumellipticum1a 0530a 12.1 132.5 Ggrovi1a 0530a 12.0 146.4 Diploneisovalis1a 0530a 13.0 147.0 Planothidiumconspicuum2a&b 0530a 13.0 138.0 Fconstruensvventer1a 0530a 14.0 127.7 Planrostratum 0530a 14.0 134.0 Planconspicua3a&b 0530a 14.0 134.7 Planconspicua4a 0530a 14.0 140.4 Fconstruv.venter2a 0530a 13.9 147.2 Fconstruv.venter3a 0530a 15.0 146.9 Fconstruens1a 0530a 15.0 140.4 Fconstruv.venter4a 0530a 15.0 138.5 Fconstruv.venter5a 0530a 15.0 137.2 Fconstruv.venter6a 0530a 15.0 136.3 Placonthidiumellipticum2a 0530a 15.0 133.2 203

Placonthidiumrostratum3a&b 0530a 15.0 131.9 Fconstruv.venter7a 0530a 15.0 129.9 Fconstruv.venter8a 0530a 15.0 128.9 Placonthidiumellipticum3a 0530a 15.0 127.5 Placonthidiumconspicuum5a&b 0530a 16.0 129.5 Diploneisovalis2a 0530a 16.0 132.1 Fconstruv.venter9a 0530a 16.0 135.7 Placonthidiumellipticum4a 0530a 13.1 129.4 NR010530 0530a 11.9 148.7 TH030530 0530a 12.8 148.4 13F.consts.v.venter0530 0530a 12.0 148.4 2Fconst.0530 0530a 12.1 148.2 1D.heimalev.mesodon 0530a 12.2 147.3 NR020530 0530a 12.3 147.1 1Fbrev.v.subcap0530 0530a 12.3 144.9 14Fconst.v.vent0530 0530a 12.2 142.9 FL010530 0530a 12.3 132.7 15F.const.v.vent 0530a 12.3 130.9 Plac.amph10530 0530a 12.3 130.3 Planothidiumconspicuum6a 0530a 16.0 138.1 Fconstru.v.venter10 0530a 16.0 138.2 Synedravaucheria1 0530a 16.0 140.5 Synedravaucheria2 0530a 17.0 141.7 Planothidiumconspicuum7 0530a 17.0 136.7 Planothidiumrostratum4 0530a 18.0 137.9 Diat.Anceps1 0526b 12.0 142.3 Nscutiloides1 0526b 12.1 140.1 Fconstru.v.venter11 0526b 12.0 134.2 Meridoncirculare1 0526b 13.0 135.6 Fconstru.v.venter12 0526b 13.0 139.2 TE010530 0530a 12.1 127.3 Fconstru.v.venter16 0530a 12.0 128.4 Fconstru.v.venter17 0530a 11.9 131.5 FL020530 0530a 13.7 135.3 Nscutelloides020530 0530a 13.6 135.6 Fconstru.v.venter18 0530a 13.6 139.0 FcuBinodis010530 0530a 13.6 144.5 Fconstru.v.venter19 0530a 13.6 145.7 Fconstru.v.venter20 0530a 13.6 147.1 Placamph020530 0530a 15.4 144.3 Fconstru.v.venter21 0530a 15.4 143.7 TH040530 0530a 15.3 138.7 CC040530 0530a 15.3 138.4 204

FnLen01 0530a 15.3 134.6 GC010530 0530a 15.4 132.4 NitzA010530 0530a 15.4 134.6 Sparas010530 0530a 15.4 135.7 GLvI010530 0530a 15.4 138.4 Placamph030530 0530a 15.3 144.3 FPvL010530 0530a 15.4 147.1 FL030530 0530a 17.0 134.4 TH050530 0530a 17.0 146.7 Fconstru.v.venter22 0530a 17.0 134.2 SS010530 0530a 17.0 130.5 CV020530 0530a 17.0 129.4 GT010530 0530a 17.0 128.9 Fconstru.v.venter23 0530a 19.0 130.2 SS020530 0530a 19.0 130.7 SS030530 0530a 19.1 131.5 02Gomphgrov0530 0530a 19.1 131.8 FnLen02 0530a 19.1 137.0 EV030530 0530a 19.0 144.1 FB010530 0530a 19.0 146.7 Tlance0537aTwillpaper 0537a 14.1 128.8 Twillgirdleband 0530a 22.2 145.1 GGvd010704b 0704b 10.2 144.2 EV040704b 0704b 10.2 141.1 FV010704b 0704b 10.2 140.9 EL010704b 0704b 10.2 139.5 FV020704b 0704b 10.2 138.0 GAvM010704b 0704b 10.2 137.8 GAvM020704b 0704b 10.2 137.6 EV050704b 0704b 10.2 137.3 GLvI020704b 0704b 10.2 137.1 GC020704b 0704b 10.2 137.1 GLvI030704b 0704b 10.3 136.9 TLac040704b 0704b 10.2 136.8 FV030704b 0704b 10.2 136.4 GC040704b 0704b 10.2 135.8 FV040704b 0704b 10.3 135.8 FV050704b 0704b 10.3 135.8 FV060704b 0704b 10.2 134.9 FV070704b 0704b 10.3 134.9 GC050704b 0704b 10.3 134.9 FV080704b 0704b 10.2 132.0 GLvI040704b 0704b 10.2 132.6 205

Tlin010707a 0707a 11.0 135.7 FV090707a 0707a 11.0 136.2 02Mcirculare0707a 0707a 11.0 136.9 EL020707a 0707a 10.9 138.6 GLvI050707a 0707a 10.8 140.0 GC060707a 0707a 10.8 140.8 GT020707a 0707a 10.7 141.7 02Diathemalv.mesodon 0707a 14.0 146.6 GT030707a 0707a 13.9 143.9 TLata010707a 0707a 13.9 136.9 03Diathemalv.mesodon 0707a 13.9 134.2 Tlin020707a 0707a 16.4 134.7 Tlin030707a 0707a 16.4 135.6 04Diathemalv.mesodon 0707a 16.3 136.2 EPvM010707a 0707a 16.4 137.5 EP020707a 0707a 16.1 138.9 PMaj010706a 0706a 16.0 134.5 PMaj020706a 0706a 16.4 139.1 SK010706a 0706a 13.1 139.1 GT040706a 0706a 13.8 139.1 CC030706a 0706a 15.7 131.5 GC070706a 0706a 15.7 138.2 206

Appendix 4 – Error Calculation Counts

Differences between slides counted per sample resulting in error. Error calculation is then listed for each sample. 207 ilis

Slides Species Actinocyclus mot Actinocyclus krasskei Actinocyclus sp Actinocyclus Anomoeoneis sphaerophora agassizi Aulacoseira Aulacoseira canadensis Aulacoseira granulata Aulacoseria granulata diagenetic Craticula sp. cistula Cymbella Cymbella ventricosa Diatoma anceps Diatoma hemalie Q0412-11 07AM05-19a 1 2 3 6 3 07AM05-20a 2 1 2 07AM05-21a 1 1 2 07AM05-22a 3 3 07AM05-23a 3 10 8 07AM05-24a 5 2 4 11 07AM05-25a 8 1 2 3 2 1 07AM05-26a 4 1 8 4 4 4 2 1 07AM05-27a 3 3 1 4 1 07AM05-28a 1 4 5 1 1 07AM05-29a uncounted, no error 07AM05-30a (3 row 387 ct) Recount necessary 07AM05-31a (++ broken up) 2 1 10 2 2 1 07AM05-32a 1 1 1 1 2 1 5 1 07AM05-33a 13 229 9 07AM05-34a (++broken up) 1 11 5 3 1 11 07AM05-35a 6 2 16123 1 07AM05-36Aa (min. presv) 2 4 121 07AM05-36Ba 6 3 4 07AM05-37a 20 32125 07AM05-38a 1 1512 11 07AM05-39a 9 1145 1 ALP 07AM07-02a 5 21115 111 07AM07-03a 5 192 121 07AM07-04a 2 2 3 41 208

07AM07-05a 4 3 3 12 07AM07-06a 2 7 6 3 07AM07-07a 2 276 2 07AM07-08a 4 2263 1 126 11111 1 11129931 21 31 2 11 111 1 Diatoma hiemale v mesodon

Diploneis ovalis 11111 11 1 1 1225 111 1121 2 1 41 1 11 41 Ellerbeckia balleyi

114151 4 1 11 11 Eunotia curvata

Eunotia pectinalis

Eunotia pseudopectinalis

Eunotia pectinalis var minor

Eunotia veneris 1 1 111 1 1 2111 510 2 Pseudostaurosira brevistriata

Pseudostaurosira brevistriata var elliptica

Pseudostaurosira brevistriata var subcapitata

52 15 Fragilaria capucina var laceolata

Staurosira construens

Staurosira construens var venter

31 3 Fragilaria lapponica

1 Fragilaria near lenoblei

Fragilaria leptostauron

3 Fragilaria pinnata var laceolata 12 1 1 Fragilaria virescens

Fragilariopsis (Fragilaria?) spVanlandingham 1964 209 210

1116 312 4 1 1 11 Gomphonema acuminatum var montanum

1111 1 3 Gomphonema angustatum

23 4 3 32 52 1111311 1 Gomphonema cholnokytes

Gomphonema gracile v. dichotoma

Gomphonema gracile v naviculoides

11 11111 2 11 Gomphonema grovei

Gomphonema grunowii 11 121 2 11 Gomphonema affine var insignious

21 Gomphonema dichotomum

1 Gomphonema tenellum (?) 1 2 12 11 1 1 Melosira undulata

Meridion circulare

Navicula acceptata

Navicula amphibola

Navicula avenacea

1 Navicula gastrum

Navicula near insula

Navicula pantoesekiama

Navicula pseudoscutiformis

Navicula reimertes 211 212

221 11 2221 1 111 11 Navicula scutelloides

43221221 24 1 2 Navicula seminuloides

Neidium iridis 1 1131 22 12 Nitzchia amphibia

121 11 1 1 11 Opephora glangeaudi

1 111 Opephora martyi

Pinnularia major

1221 Pinnularia nodosa

Planothidium conspicuum

Planothidium ellipticum

Planothidium lanceolatum

Planothidium rostratum

Rhoicosphenia curvata

Sellophora bacillum

Stauroneis kriegeri

Synedra tabulata var fasiculata

1 Synedra near socia (my diagnosis)

Synedra vaucheriae

1 Synedra sp. (close to fragilariopsis ?) 1 1 Tetracyclus ellipticus 213 214

1 11 11 11 1 2%(AG)and0.5% 3%to1% 1 1%to0.3% 1 2 1 1 1 2 1 0.50% 1 1 Tetracyclus lacustris 1 1 211 211 4%(EB)and1% 1 1 1 Tetracyclus lata 12 11 3%(AG)and0.5% 1 Tetracyclus lancea

1 Tetracyclus linearis 4%(Acti)6%(AG)and0.5% 1 1 3%and1% 1 Tetracyclus polygibbum

Tetracyclus sp H

Tetracyclus sp H lanceate

Unknown sp (with photos) recount no counti.e.error 3% (Acti)and0.5% 4% (AG)and1% 3% (Aulac)and0.5% 1% 0.50% ~1% 3% to0.5% 5% (AG)and1% 3% (Acti)and1% 3% (AG)and2-0.5% 7% (Acti)and4%(AC)&0.5% 2% to0.5% 1% to0.5% 5% (AG)and1% 2% to0.5%

Calculated Error 215 216

11 2% to 0.5% 1 2% (Aulac) and 0.5% 3 2% (Aulac) and 0.5% 11 2% (AC) and 1% 217

Appendix 5 – Measurements of Width for Grain Size Calculation

Grain size measurements of samples chosen to represent each unit within the diatomite. Measurements are of the width/diameter dimension. Mean and mode measurements for each sample are listed. 218

Slide counted Subunit Specimen Width 07AM05-20a TC 6 07AM05-20a TC 8 07AM05-20a TC 9 07AM05-20a TC 5 07AM05-20a TC 8 07AM05-20a TC 7 07AM05-20a TC 7 07AM05-20a TC 12 07AM05-20a TC 6 07AM05-20a TC 7 07AM05-20a TC 5 07AM05-20a TC 7 07AM05-20a TC 7 07AM05-20a TC 6 07AM05-20a TC 7 07AM05-20a TC 7 07AM05-20a TC 9 07AM05-20a TC 4 07AM05-20a TC 5 07AM05-20a TC 7 07AM05-20a TC 9 07AM05-20a TC 5 07AM05-20a TC 10 07AM05-20a TC 7 07AM05-20a TC 5 07AM05-20a TC 8 07AM05-20a TC 5 07AM05-20a TC 5 07AM05-20a TC 4 07AM05-20a TC 6 07AM05-20a TC 7 07AM05-20a TC 7 07AM05-20a TC 10 07AM05-20a TC 9 07AM05-20a TC 8 07AM05-20a TC 10 07AM05-20a TC 6 07AM05-20a TC 7 07AM05-20a TC 6 07AM05-20a TC 7 07AM05-20a TC 8 07AM05-20a TC 7 07AM05-20a TC 6 07AM05-20a TC 7 07AM05-20a TC 5 07AM05-20a TC 6 07AM05-20a TC 7 219

Avg: 6.96 Mode 7 220

07AM05-20a TC 6 07AM05-20a TC 9 07AM05-20a TC 7 07AM05-25a TC 52 07AM05-25a TC 8 07AM05-25a TC 10 07AM05-25a TC 7 07AM05-25a TC 9 07AM05-25a TC 9 07AM05-25a TC 6 07AM05-25a TC 8 07AM05-25a TC 12 07AM05-25a TC 8 07AM05-25a TC 6 07AM05-25a TC 5 07AM05-25a TC 7 07AM05-25a TC 7 07AM05-25a TC 7 07AM05-25a TC 8 07AM05-25a TC 46 07AM05-25a TC 11 07AM05-25a TC 26 07AM05-25a TC 7 07AM05-25a TC 8 07AM05-25a TC 47 07AM05-25a TC 7 07AM05-25a TC 12 07AM05-25a TC 9 07AM05-25a TC 38 07AM05-25a TC 6 07AM05-25a TC 20 07AM05-25a TC 24 07AM05-25a TC 7 07AM05-25a TC 10 07AM05-25a TC 10 07AM05-25a TC 42 07AM05-25a TC 8 07AM05-25a TC 7 07AM05-25a TC 7 07AM05-25a TC 5 07AM05-25a TC 46 07AM05-25a TC 10 07AM05-25a TC 10 07AM05-25a TC 10 07AM05-25a TC 6 07AM05-25a TC 17 07AM05-25a TC 8 07AM05-25a TC 46 07AM05-25a TC 9 07AM05-25a TC 10 221

Avg: 15.68 Mode: 7 222

07AM05-25a TC 39 07AM05-25a TC 38 07AM05-25a TC 9 07AM05-30 BC 6 07AM05-30 BC 4 07AM05-30 BC 3 07AM05-30 BC 4 07AM05-30 BC 6 07AM05-30 BC 5 07AM05-30 BC 5 07AM05-30 BC 4 07AM05-30 BC 5 07AM05-30 BC 4 07AM05-30 BC 12 07AM05-30 BC 8 07AM05-30 BC 3 07AM05-30 BC 5 07AM05-30 BC 9 07AM05-30 BC 4 07AM05-30 BC 5 07AM05-30 BC 8 07AM05-30 BC 5 07AM05-30 BC 5 07AM05-30 BC 5 07AM05-30 BC 8 07AM05-30 BC 3 07AM05-30 BC 3 07AM05-30 BC 24 07AM05-30 BC 3 07AM05-30 BC 5 07AM05-30 BC 4 07AM05-30 BC 7 07AM05-30 BC 5 07AM05-30 BC 4 07AM05-30 BC 3 07AM05-30 BC 16 07AM05-30 BC 4 07AM05-30 BC 3 07AM05-30 BC 4 07AM05-30 BC 7 07AM05-30 BC 5 07AM05-30 BC 8 07AM05-30 BC 44 07AM05-30 BC 45 07AM05-30 BC 62 07AM05-30 BC 61 07AM05-30 BC 9 07AM05-30 BC 17 07AM05-30 BC 42 07AM05-30 BC 64 223

Avg: 14.92 Mode 5 224

07AM05-30 BC 59 07AM05-30 BC 58 07AM05-30 BC 54 07AM05-37a 4C 7 07AM05-37a 4C 7 07AM05-37a 4C 22 07AM05-37a 4C 32 07AM05-37a 4C 9 07AM05-37a 4C 7 07AM05-37a 4C 22 07AM05-37a 4C 4 07AM05-37a 4C 11 07AM05-37a 4C 8 07AM05-37a 4C 10 07AM05-37a 4C 7 07AM05-37a 4C 6 07AM05-37a 4C 17 07AM05-37a 4C 6 07AM05-37a 4C 4 07AM05-37a 4C 6 07AM05-37a 4C 6 07AM05-37a 4C 8 07AM05-37a 4C 7 07AM05-37a 4C 7 07AM05-37a 4C 7 07AM05-37a 4C 7 07AM05-37a 4C 8 07AM05-37a 4C 6 07AM05-37a 4C 4 07AM05-37a 4C 6 07AM05-37a 4C 5 07AM05-37a 4C 11 07AM05-37a 4C 11 07AM05-37a 4C 5 07AM05-37a 4C 8 07AM05-37a 4C 19 07AM05-37a 4C 16 07AM05-37a 4C 9 07AM05-37a 4C 4 07AM05-37a 4C 11 07AM05-37a 4C 4 07AM05-37a 4C 35 07AM05-37a 4C 12 07AM05-37a 4C 4 07AM05-37a 4C 6 07AM05-37a 4C 6 07AM05-37a 4C 5 07AM05-37a 4C 6 07AM05-37a 4C 6 07AM05-37a 4C 7 225

Avg: 9.22 Mode 6 226

07AM05-37a 4C 5 07AM05-37a 4C 10 07AM05-37a 4C 5 227

Appendix 6 – Extended Methodologies

Extended methodology not provided in Chapter 1. Extended methods include specific cores sampled and preparation techniques. 228

Extended Methodologies

Sampling

Data was analyzed from four localities; World Minerals mining locality drill cores were sampled from the Frenchman Hills and the northern mining sections near the Gorge

Amphitheater (approximately 5 km north of the Frenchman Hills), the Quincy Ancient

Lakes Park in the Ancient Lakes Trail Campground, and from the United States Army

Core of Engineers drilling cores on the North end of Moses Lake and Potholes Reservoir

(Figure 1a, Table 1). However, the Moses Lake cores were extensively altered and did not produce diatoms, therefore only the first three localities are discussed in detail for this study.

All samples were collected in 3 inch by 6 inch plastic sample bags. These sample bags have zipper style seals and were made of heavy-grade plastic so as to prevent contamination. All samples were placed in a separate sample container after collection from outcrop or core to prevent contamination while collecting in the field.

The Quincy Diatomite is best exposed in mining locations owned and operated by

World Minerals Inc. However, World Minerals actively mines only the Top Crude leaving the remaining subunits unexposed in mining trenches. As a result, the best exposures of complete sections come from exploratory drill cores done by World

Minerals. These drill cores represent stratigraphically complete sections from the top to the bottom of the Quincy Diatomite. World Minerals stores these cores in the town of

Quincy, WA at the Celite® Diatomite Products plant and granted access to both the stored drill cores, and their active mining locations. 229

Samples were procured from three stratigraphically complete World Minerals drill cores. All samples and their corresponding cores appear in Appendix 1. Core

Q0412-11 comes from a section recently active in the Frenchman Hills (Figure 1a, b).

This core was sampled from the base of the diatomite (50.8 ft depth) to its top (16.8ft depth). Twenty-three samples were taken from this core at an approximate interval of 30 cm, with some minor variation in sampling interval depending on core recovery, transition between diatomite units, and observation of lithologic variation, which were sampled more closely. Position of samples from the core appears in figure 3.

Core Q01-20-9 comes from the second locality, an actively mined section near the

Gorge Amphitheater, approximately 5 kilometers north of the Frenchman Hills (Figure

1a). This core was also sampled from the base of the diatomite (23ft depth) to the upper contact with the overlying basalt (15 ft depth). Six samples were collected from this core at an interval of approximately 1.5 ft (Figure 3). This core only contained the Top Crude; sample positions from the core appear in figure 3. Samples from this core were examined but not counted.

The third locality is an outcrop at the Quincy Ancient Lakes Park (ALP). The park is approximately 10 kilometers north of the Frenchman Hills and is part of the

Ancient Lakes Trail Campground. This location has an isolated outcrop of the Top

Crude exposed on the south-facing hillside along the northern main trail (Figure 1a, c).

The outcrop exposes 3 m of section of Top Crude, with the base of the diatomite being covered by soil development and thick vegetation on the hill. It is unclear whether the

Four Crude or the Bottom Crude are present at the ALP due to the limited exposure, though the small area left unsampled (approximately 2 meters) suggests only the Top 230

Crude occurs at this locality. Nine samples were collected from this locality at 0.5-meter intervals over the course of exposed section, and these samples correlate with the Top

Crude unit (Figure 3).

Locality four is approximately 35 miles east of the Frenchman Hills on the northern shore of Moses Lake (Figure 1a). The United States Army Core of Engineers

(USACE) has done exploratory drilling projects around Larson Air Force base, and has encountered the Quincy Diatomite deposit at depth. The USACE granted access to these cores, and samples were collected at 0.3-meter intervals, as the deposit is reduced to less than 1 meter thickness at this location. Eight samples were taken from three cores;

00BW14 (three samples from 74-76ft), 00BW06 (two samples from 175-177.5ft), and

99BW15 (two samples from 98-99ft). Samples from these cores are correlated to the Top

Crude. All samples from this locality were significantly more indurated and thermally altered than samples from the three localities to the west, and diatoms were not preserved

(only a single poor fragment was observed in light microscopy).

Lab Work

All samples were cleaned by placing approximately 1-3 grams of diatomite in 1 dram glass vials. Vials were filled with 3 percent, home pharmaceutical grade hydrogen peroxide solution until the peroxide just covered the dry samples. The vials were then topped off with distilled water. Samples were allowed to settle for at least 8 hours and were then decanted and refilled with distilled water 6 times to remove any residual clay particles.

A diluted slurry was derived from the washed sample vials by using several drops of the original slurry in a separate shell vial, then adding distilled water. Distilled water 231 was added until the solution reached the “eye-clear” point when the slurry was no longer cloudy and particles were visible only on close inspection. This diluted slurry was used to coat 0.1mm thick cover slips, distributing the material evenly. Cover slips were then allowed several hours on a vibration-free surface under a low intensity heat lamp until dry, and were then mounted onto 1.0mm thick, standard width glass slides.

Mounting on permanent slides was done using either Zrax© mounting medium

(refractive index ~1.7) or Piccolyte (refractive index at least 1.52). Mounting medium was placed on the slide and the coated cover slip was placed on top. The slides were then put on a hotplate set to 225oC for several minutes allowing the mounting medium to devolatilize. Slides were removed from the hotplate when the degassing appeared to be complete and the mounting medium became visibly more viscous. The slides were then allowed to cool and the mountant hardened.

At least two slides of each sample were made (yielding over 200 slides total).

Slides were examined using an Olympus BX-51 microscope equipped with a 100x oil immersion lens, DIC optics, and 1.4NA. Identification of species present in each sample was done at 400X and 1000X magnification and light micrographs were taken using an

Olympus DX-71 microscope camera with Olympus software. Coordinates of specimens photographed using light microscopy are provided in Appendix 7.

Specimens were also examined and identified using a liquid nitrogen cooled Jeol

JSM 840A Scanning Electron Microscope equipped with Fissions Kevex digital beam control interface and software for photography. Samples were examined at up to

100,000x magnification for feature identification and digital imaging. SEM stubs were coated with the residual diluted slurry material from permanent slide preparation and 232 allowed to air-dry. Sample stubs were sputter coated with gold for at least 2 minutes per stub to ensure good picture resolution and minimal “charging” while viewing under the

SEM.

Frustule point counts were done in a linear transect on each slide for cores Q0412-

11 and for samples from the ALP. The counting transect was started at the Y = 12.0 marker on the microscope stage, at the extreme left of the coverslip. Frustules were then counted moving left to right along the Y = 12.0 line until 150 frustules had been counted

(yielding counts of at least 300 per sample). If fewer than 150 frustules were counted and across the one line, the slide was moved to Y = 13.0 to resume a transect count, then

14.0, and so on until the count was complete. Frustules were only counted if they were greater than 50% intact. Point count data is presented in Appendix 2. Error calculations for point count data were made by comparing count differences between slides within samples, and calculating the percent error of the species counted. The differences in relative abundance were then calculated and compared to the overall counts to get the range of error for all samples. Error calculation data is presented in Appendix 3.

Four samples representing the various units within the stratigraphy were examined and fifty complete frustules were measured along their width. The measurements were taken starting from the top of the slide at the X = 145.0 mark, and moving down along the Y axis until the 50 frustules had been counted. These measurements were then used to determine the mean and mode grain size of each unit, with the mean and the mode grain sizes being plotted in Appendix 4.

Error within Relative Abundance Counts 233

Error for the relative abundance counts shows minimal variance across all samples and localities. Comparison of the two slide counts per sample yielded an error range of 0.3% to 7% among all samples, with most samples error being between 0.5% and 2% (Appendix 3). The highest errors for these samples were principally from counts of Aulacoseira granulata, which was counted most frequently and thus most subject to error in counts. One sample in the Frenchman Hills (sample 07AM05-37, depth 8.8m) displayed the highest error of all, with a 7% error in counting Actinocyclus motilis

(Appendix 3). This sample is part of the Four Crude.

Samples from core Q0412-11 and the ALP were also processed and run to test for any remaining organic carbon, with a total of 10 samples being run, incorporating the entire stratigraphy from each location. Approximately 1 gram of 7 samples from the

Frenchman Hills and 3 samples from the ALP were placed in 5ml glass vials. These samples were then placed in an acid fume bath for 1 week in order to dissolve any trace carbonate material that may be present. Samples were then removed, powdered, and put into aluminum boats for processing conducted by Dr. Simon Poulson, to measure any

13 released CO2 (via C) from incinerated organic mater.

Data Analysis

Point count data were analyzed using the program R, version 2.9.1, available in the public domain. Extremely rare taxa (those with less than 2% total occurrence in any sample, and those with less than two occurrences over all samples counted) were removed from the data set in order to create a more parsimonious model. NonMetric

Multidimensional Scaling (nMDS) was used to analyze counts from the Frenchman Hills and the Ancient Lakes Park. NonMetric Multidimensional Scaling is an unconstrained 234 ordination useful as an alternative to Correspondence Analysis. Matrix calculations used in this technique are less sensitive to non-linear relationships and discontinuous distributions, making the method more likely to display the real differences between species and samples and provide more robust results for interpretation of data (Clarke and

Ainsworth 1993, McCune and Grace 2002). Samples within the dataset were given the sample numbers used during collection, and species were identified using letters starting with D and continuing through to double letter denotation (Table 2). Data from nMDS calculations were plotted against factors 1 through 3 for analysis of which species and samples group together and how the species and samples vary between each other.

NMDS plots are representations of multidimensional space in a two dimensional plot. Axes of the plots represent ecologic gradients (in the case of Miocene material, the specific ecologic conditions these gradients represent are often recondite). Species plot in arrangement of their distribution across all samples; species near the origin are common to all samples, while those species plotting far from the origin and near specific samples show a high correlation with those samples. Species and samples plotting near each other on the graph are considered statistically related.

Arrows for samples represent vectors, showing the degree of influence any given sample has on the overall data, and how unique each sample is in terms of floral assemblage. Sample vectors with the longest arrows contain the most distinctive floral assemblages, while those samples plotting near the origin with shorter arrows share many species in common, and are not considered highly influential to the dataset. Arrows at

900 from each other show no association, while those arrows at 1800 have an inverse 235 relationship. Samples plotting near the origin share many common species and have floral assemblages likely to resemble other samples.

Samples were also analyzed with a cluster analysis using Wards minimum variance method for determination of relatedness among the samples. Wards method of clustering is an agglomerative clustering technique that clusters samples based on

Euclidean distances (absolute and relative) between species and samples; the clusters are progressively expanded based on minimum variance of within-group total sum of squares. This cluster analysis preserves the “original distance” between samples, and is interpreted to show a more natural ecologic clustering of data samples based on species composition (McCune and Grace 2002). The results of this cluster analysis were then compared to those from nMDS for interpretation of relatedness of samples and taxa.

236

Appendix 7 – Organic Carbon Test Results

Results of test for organic carbon 13C. Samples are listed by number and unit sampled. Sample 07AM07-09 corresponds to 07AM07-02 at the top of the ALP section 237

Results of test for organic carbon 13C. Samples are listed by number and unit sampled. Sample 07AM07-09 corresponds to 07AM07-02 at the top of the ALP section.

Sample # Unit Sampled Location 13C (‰ vs. VPDB) Wt. % C Frenchman Hills 07AM05-20 Top Crude FH -38.6 0.73 07AM05-23 Top Crude FH -35.8 0.51 07AM05-25 Top Crude FH -39.1 0.96 07AM05-28 Bottom Crude FH -37.6 1.14 07AM05-32 Bottom Crude FH -38.2 0.97 07AM05-35 Four Crude FH -41.0 1.89 07AM05-38 Four Crude FH -38.8 0.84 Mean Wt. %C =1.0 ALP 07AM07-09 Top Crude ALP -39.7 0.79 07AM07-04 Top Crude ALP -37.5 0.98 07AM07-07 Top Crude ALP -37.0 1.15 Mean Wt. %C =0.97 238

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