Paleo-upwelling and the distribution of Mesozoic marine reptiles

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PALEO-UPWELLnSfG AND THE DISTRIBUTION OF MESOZOIC

MARINE REPTILES

by

Danielle Dawn Montague-Judd

Copyright © Danielle Dawn Montague-Judd 1999

A Dissertation Submitted to the Faculty of the

DEPARTMENT OF GEOSCIENCES

In Partial Fulfillment of the Requirements For the Degree of

DOCTOR OF PHILOSOPHY

In the Graduate College

THE UNIVERSITY OF ARIZONA

1999 XJMI Niimber: 9946864

Copyright 1999 by Montague-Judd, Danielle Dawn

All rights reserved.

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THE UNIVERSITY OF ARIZONA ® GRADUATE COLLEGE

As members of the Final Examination Committee, we certify that we have

read the dissertation prepared by Danielle Dawn Montague-Judd

entitled Paleo-upwelling and the Distribution of Mesozoic Marine Reptiles

and recommend that it be accepted as fulfilling the dissertation requirement for the Degree of Doctor of Philosophy

jlO j'i ^ idith T. Parrish-Jones Date ^ // a /py Date

—

Date '

Philip A. Hastings Date

Final approval and acceptance of this dissertation is contingent upon the candidate's submission of the final copy of the dissertation to the Graduate College.

I hereby certify that I have read this dissertation prepared under my direction and recommend that it be accepted as fulfilling the dissertation requirement.

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STATEMENT BY AUTHOR

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ACKNOWLEDGMENTS

Many people willingly gave of their time and expertise to help me complete this dissertation. Judy Parrish, my major advisor, introduced me to the topic, helped instill in me the confidence to proceed, and provided academic support. Karl Flessa acted as my interim advisor during Judy's sabbatical and included me in the C.E.A.M. community. Andy Cohen, John Lundberg, and Phil Hastings provided guidance and helpful advice. My whole committee deserves thanks for promptly reviewing the dissertation. Tim Demko, Bob Moulton, and Amy Kowalczyk served excellently as field assistant-mentors, and Amy Kowalczyk and Essa Gross prepared rock samples. Wes Bilodeau and Joe Schreiber gave expert advice on petrography and access to petrographic equipment. Mark Rigali and Pete Holterhoff gave me crash courses on thin-section petrography that helped immensely. Mark Rigali, Joaquin Ruiz, and Mark Baker provided advice on geochemical analysis. Pete Holterhoff and Bob Casavant gave stimulating discussion of carbonate sedimentology. Kate Rylander, Andy Cohen, and Pete DeCelles allowed use of microscopes and video-imaging equipment. Kevin Moodie provided fossil preparation equipment and methods. Julie Libarkin helped me understand Luning paleomagnetic data. Thanks to the 1993 and 1994 University of Arizona field camp directors (J. Parrish, S. Beck) and staff for allowing me to tag along. Jennifer Hogler oriented me to the Luning Formation at BISP. Norm Silberling graciously provided conodont information and insight on rocks of the MMP. Chris McRoberts identified bivalve fossils from BISP. George Gehrels helped me get the big picture of Mesozoic accretionary tectonics and substantially increased my understanding of the Luning paleogeographic setting by giving me passage on the DUDZ tour of Nevada and California in 1995. Jeff Manuszak provided the Luning detrital zircon data. Big thanks to Tom Moore for giving me access to his geodesic software, for many stimulating discussions, and for help in many aspects of the database study. Thanks to Karen Porter, Aaron Bell, Dave Dettman, and Tong Jinnan for article translations. Chris Scotese generously gave paleogeographic maps and software. A.M. Ziegler and D. Rowley of the Paleogeographic Atlas Project, University of Chicago, graciously provided upwelling-related lithology data. Barry Kazmer dug up helpftil taxonomic information. Among the many who helped me complete this project through discussion or support at various times are Bill Dickinson, Peter Coney, George Davis, Judy Massare, George Stanley, Martin Sander, Mike Everhart, Pete DeCelles, Conrad Labandeira, Lois Roe, Dena Smith, Tara Curtin, Julie Libarkin, Kim Driver, Peter Roopnarine, and the PCRP lab group. Pat and Fred Montague inspired me to pursue a science career. Curtis Judd provided multi-faceted and untiring support—computing, field, emotional, mental. Finally, this project would not have happened without monetary support from the National Science Foundation, Phi Kappa Phi, the University of Arizona Graduate College, the Geological Society of America, the American Association of Petroleum Geologists, Sigma Xi, the University of Arizona Research Training Group in the Analysis of Biological Diversification, and the Marshall Foundation. 5

TABLE OF CONTENTS

List of Figures 8 List of Tables 9 ABSTRACT 10 1. INTRODUCTION AND OVERVIEW OF MODERN AND ANCIENT UP WELLING 12 INTRODUCTION 13 MODERN UPWELING SYSTEMS 15 Physical Basis 19 Biological Responses to Upwelline 19 Nutrients 19 Plankton 21 Nekton 22 Benthos 25 Biological Responses: Summary 30 Sedimentological Consequences of Upwelling 31 General Patterns 31 Organic-Rich Sediments 33 Biosiliceous Sediments ("Chert") 37 Phosphorite 38 Glauconite 40 Calcareous Ooze/Chalk 40 Trace and Minor Element Concentrations 42 Sedimentological Consequences: Summary 43 Sources of Variable Sedimentation 44 ANCIENT UPWELLING SYSTEMS 48 Preservable Aspects of Upwelling Sediments and Biota 48 Evaluation 50 Examples of Ancient Upwelling Systems 53 SUMMARY 59 2. PALEOCEANOGRAPHY OF THE UPPER TRIASSIC LUNING FORMATION AT THE SHOSHONE MOUNTAINS, NEVADA 71 INTRODUCTION 72 TECTONOSTRATIGRAPHIC SETTING 77 LITHOLOGY, AGE, AND PALEONTOLOGY OF THE LUNING FORMATION 82 METHODS 89 RESULTS 91 Lithofacies Descriptions and Interpretations 91 Shaly. Fossiliferous Limestone Facies (S) 92 Description 92 6

TABLE OF CONTENTS - Continued

Siltv. brown limestone and argillite subfacies fSla, Figure 2.7). 92 Siltv. black limestone subfacies (Sbl. Figure 2.8). 94 Calcareous shale subfacies fScs. Figure 2.9). 96 Interpretation 97 Pel-Bioclastic Limestone Facies fP) 102 Description 102 Peloidal bioclastic limestone subfacies (Pel. Figure 2.10). 102 Pelleted bioclastic limestone subfacies fPbl. Figure 2.11). 103 Interpretation 104 Oolitic Bioclastic Limestone Facies (O. Figure 2.12) 105 Description 105 Interpretation 106 Fossiliferous Limestone Facies (F. Figure 2.13) 106 Description 106 Interpretation 107 Geochemical Results 108 Paleontological Results 109 DISCUSSION Ill Environments of Deposition Ill Sedimentological Aspects 111 Geochemical Aspects 113 Paleontological Aspects 116 Upwelling Characteristics 117 Cool-Water Conditions 117 Edge Effects 119 Eutrophic Conditions 120 Synthesis 122 Scenario 1 123 Scenario 2 124 Scenario 3 126 CONCLUSIONS 129 3. PALEOUPWELLING AND THE DISTRIBUTION OF MESOZOIC MARINE REPTILES 162 INTRODUCTION 163 Modem Whale Distribution and Feeding 163 Marine Reptile Ecology 165 Upwelling Characteristics 167 Overview of the Studv 167 METHODS 169 Database Design and Assembly 169 Upwelling-Associated Lithologies 171 7

TABLE OF CONTENTS - Continued

Paleogeographic Reconstructions and Upwelline Predictions 172 Data Plotting Procedures 174 Geographic Analysis of Gridded Data 175 Distance Analysis of Gridded Data 176 RESULTS 180 Database and Data Processing 180 Geographic Analysis of Gridded Data 181 Distance Analysis of Gridded Data 183 DISCUSSION 186 Sampling Biases 187 Preservational Biases 189 Analytical Biases 192 Marine Reptile Ecology 195 CONCLUSIONS 201 APPENDICES A. MEASURED STRATIGRAPHIC SECTIONS 227 B. RESULTS OF INORGANIC AND TOC ANALYSES FOR SAMPLES FROM THE LUNING FORMATION AT WEST UNION CANYON 260 C. DATA USED FOR FIGURE 2.16 268 D. MARINE REPTILE DATABASE TABLE RELATIONSHIPS, STRUCTURES, AND FIELD CODES 277 E. TAXONOMIC SCHEMES USED IN THE MARINE REPTILE DATABASE 285 F. PALEGEOGRAPHIC BASE MAPS AND UPWELLING PREDICTIONS 295 G. MARINE REPTILE DATABASE 301 H. DATABASE REFERENCES 354 I. GRIDDED AND RAW DATA FOR REPTILES AND LITHOLOGIES, PLOTTED ON PALEGEOGRAPHIC BASE MAPS 365 J. DISTANCE CALCULATION RESULTS 408 REFERENCES 420 8

LIST OF FIGURES

FIGURE 1.1, Simplified view of a coastal upwelling system 60 FIGURE 1.2, Today's major upwelling areas 61 FIGURE 1.3, Global phosphorite and guano island distribution 62 FIGURE 2.1, Tectonic setting of westem North America in the Late Triassic 131 FIGURE 2.2, Outcrop map of the southern Mesozoic marine province 132 FIGURE 2.3, Lithologic variation among Luning Formation localities 133 FIGURE 2.4, Geologic map of the Luning Formation in West Union Canyon 134 FIGURE 2.5, Legend and composite stratigraphic column of the Luning Formation at West Union Canyon 135 FIGURE 2.6, Correlation among stratigraphic sections in the marine part of the Luning Formation at West Union Canyon 137 FIGURE 2.7, Representative samples of the Sla subfacies 138 FIGURE 2.8, Representative samples of the Sbl subfacies 139 FIGURE 2.9, Representative samples of the Scs subfacies 140 FIGURE 2.10, Representative samples of the Pel subfacies 141 FIGURE 2.11, Representative samples of the Pbl subfacies 142 FIGURE 2.12, Representative samples of the O facies 143 FIGURE 2.13, Representative samples of the F facies 144 FIGURE 2.14, Log TOC and major element concentrations by facies 145 FIGURE 2.15, Log TOC and minor element concentrations by facies 146 FIGURE 2.16, Trends in fossil abundance (from Hogler, 1992a) 147 FIGURE 2.17, Lithofacies plotted by water depth 148 FIGURE 2.18, Ammonoid morphotype abundances by lithfacies 149 FIGURE 2.19, Scenarios for deposition of the Luning Formation 150 FIGURE 3.1, Number of localities and taxon-localities 203 FIGURE 3.2, Number of marine reptile and lithology localities 205 9

LIST OF TABLES

TABLE LI, Representative references on different aspects of upwelling 63 TABLE 1.2, Physical oceanographic characteristics of upwelling areas 64 TABLE 1.3, Primary productivity estimates for shelf regions 65 TABLE 1.4, Geologic features of modem upwelling areas 66 TABLE 1.5, Upwelling conditions and preservable upwelling indicators 67 TABLE 1.6, Comparison of four ancient upwelling deposits 68 TABLE 1.7, Suggested criteria for identifying ancient upwelling zones 70 TABLE 2.1, Preservable macrofaunal characteristics of eutrophic ecosystems 151 TABLE 2.2, Estimates of sedimentation rate for ammonoid Zones 152 TABLE 2.3, Summary of lithofacies, Luning Formation, West Union Canyon 153 TABLE 2.4, Orientations of 441 Halobia sp. shells 154 TABLE 2.5, Geochemical analysis results grouped by lithofacies 156 TABLE 2.6, Spearman rank correlation coefficients 158 TABLE 2.7, Geochemical comparison of Luning samples with other sediments....159 TABLE 2.8, Expected characteristics of the scenarios for Luning deposition 160 TABLE 2.9, Distribution of marine vertebrates in the MMP 161 TABLE 3.1, Stages of paleogeographic maps 207 TABLE 3.2, Codes and criteria used to assign data points to map intervals 208 TABLE 3.3, Calculated shelf and upwelling areas 210 TABLE 3.4, Number of localities, taxon-localities, and families 211 TABLE 3.5, Number of marine reptile and upwelling-related lithology localities ..212 TABLE 3.6, Rotational minimization results 213 TABLE 3.7, Comparison of reptile fossil occurrences and predicted upwelling 214 TABLE 3.8, Counts of grid cells containing marine reptile families 215 TABLE 3.9, Comparison of reptile family occurrences and upwelling 217 TABLE 3.10, Descriptive statistics for shortest distances, lithology types 219 TABLE 3.11, Descriptive statistics for shortest distances, reptile orders 221 TABLE 3.12, Number of grid cells containing lithologies and reptiles 223 TABLE 3.13, Lithology and taxonomic breakdown of grid cells 224 TABLE 3.14, Taphonomic states recorded for taxon-localities 225 TABLE 3.15, Counts of grid cells containing upwelling-related lithologies 226 10

ABSTRACT

Marine upwelling occurs when surface currents diverge or are deflected. Deeper

water, often nutrient-rich, rises and generates a cascade of biological effects including

elevated productivity and a unique assemblage of organisms. Macrofaunal characteristics

of upwelling provide key evidence for oxygen-minimum zones, upwelling of cool water,

and high productivity and are potentially useful indicators of ancient upwelling.

The Upper Triassic Luning Formation in Nevada contains abundant, large

ichthyosaurs and was deposited in a back-arc basin that could have experienced

upwelling conditions. Luning Formation rocks at West Union Canyon were analyzed for

sedimentological, geochemical, and paleontological upwelling indicators. Abundant

suspension feeders, lack of corals and calcareous algae, modest total organic carbon and

minor element concentrations in deeper marine facies, abundant cosmopolitan molluscs

but no taxa restricted to low latitudes, and abundant fecal pellets and clotted fabrics in

most facies suggest that upwelling could have influenced Luning deposition. Moderate- scale upwelling likely contributed to eutrophic conditions and ichthyosaur abundance at

West Union Canyon.

Marine reptiles might have had ties to upwelling areas to provide food, as do

modem whales. A relational database containing 817 locality records and 1365 taxon-

localities was assembled for ichthyosaurs, plesiosaurs, and mosasaurs. Marine reptile localities were compared with model-predicted upwelling and with upwelling-related lithologies (organic-rich rock, biogenic silica, phosphorite, and glauconite). Marine reptile occurrences intersected predicted upwelling more often than expected by chance 11 for the Upper Cretaceous, Callovian, and Norian stages, and for all of the data together {P

= 0.05). For age-restricted data, occurrences of Mosasauridae, Pliosauridae, and

Plesioauria intersected upwelling more often than expected by chance {P = 0.05).

Average shortest distances between reptile fossil and upwelling lithology occurrences were smallest (one grid cell adjacent or smaller) for the Pliensbachian and four of five

Cretaceous stages. Analytical biases and other aspects of reptile ecology may have affected the results, but overall, upwelling could have influenced marine reptile distribution, particularly for the Upper Cretaceous. Multiple radiations into the high- productivity, top-predator niche over the Mesozoic are suggested by the dominance of different taxa in grid cells containing upwelling lithologies: ichthyosaurs (early

Mesozoic), plesiosaurs (middle Mesozoic), and finally mosasaurs (late Mesozoic). 12

CHAPTER 1

INTRODUCTION AND OVERVIEW OF MODERN AND ANCIENT UPWELLING 13

INTRODUCTION

Marine upweiling occurs when surface currents either are deflected along a coast

or diverge in the open ocean and allow deeper water to surface. This deeper water is

often nutrient-rich and creates higher-than-normal biological production. Elevated

production often supports natural resources such as fisheries (Gushing 1978; Pauly and

Christensen 1995); when upweiling sediments are preserved, reserves of petroleum,

diatomite, and phosphorite can form (Brongersma-Sanders 1948; Diester-Haass 1978;

Summerhayes et al. 1995). Parrish (1995) estimated that as much as 93% of Phanerozoic oil-prone, organic-carbon-rich deposits and 82% of known phosphate deposits can be explained by upweiling in ancient seas.

Although much work has been done on the sedimentology, geochemistry, and

micropaleontology of ancient upweiling zones (Table 1.1), relatively little work has been done on the macro faunas of ancient upweiling zones (exceptions include Allmon 1993;

Allmon et al. 1996). Macrofaunal characteristics of upweiling zones are potentially

useful in identifying upweiling in the geologic record, especially where geochemical or micropaleontological data are not preserved.

In this thesis, I explore the relationship between macrofauna and ancient upweiling in two ways. First, I examine the geology and paleontology of the Upper

Triassic Luning Formation of Nevada, a possible upweiling zone deposit (Hogler 1992b).

This formation contains abundant, large ichthyosaurs and macroinvertebrates and was deposited on the western edge of North America during the Triassic, in an area that would be expected to experience coastal upweiling. I discuss the results of this analysis 14

in Chapter 2. Second, I examine the distribution of extinct marine reptiles relative to

upwelling deposits and predicted upwelling zones. Marine reptiles have been proposed

as ecological equivalents of modem whales (Parrish and Parrish 1983), which are largely

dependent on upwelling and whose evolution has been linked to tectonic and climatic

events that might have increased upwelling (Fordyce 1980; 1991). In Chapter 3 I discuss

the results of a database analysis of Mesozoic marine reptile fossils in relation to ancient

upwelling.

In this chapter, I review the physical, biological, sedimentological, and

geochemical characteristics of areas that experience upwelling today. Then, 1 review

how these systems might be preserved in the geologic record. In order to understand ancient upwelling-zone geology and paleontology, one must understand the sedimentology, chemistry, and biology of modem upwelling zones. This chapter

highlights the importance of organisms of all sizes as ecosystem components and as

indicators of ancient upwelling. Virtually all of the unique geochemical and sedimentological aspects of upwelling zones result from the biological response to increased nutrient concentrations. Currently, the geochemical and micropaleontological aspects of upwelling in the Recent are fairly well-known, meaning that ancient upwelling deposits with similar preserved characteristics can be readily identified. However, the macrofaunal characteristics of Recent upwelling are less well-known, meaning that the influence of upwelling on deposits with only macrofauna preserved is not as readily determined. Knowing how macrofauna respond to high productivity conditions can potentially broaden our understanding of ancient upwelling. 15

MODERN UPWELLING SYSTEMS

Physical Basis

Most upwelling in the oceans occurs where wind-driven surface currents diverge or are deflected off of a coastline, allowing deeper, usually cooler and more nutrient-rich water to surface (Gargett 1991; Mann and Lazier 1991). Topography and density-driven currents also can induce upwelling, but wind-driven systems today comprise the largest, most persistent, and most likely preservable upwelling environments and are the focus of this chapter. Wind-driven upwelling occurs today in both coastal and open-ocean equatorial settings and represents a mesoscale response of the ocean to large-scale winds

(Barber and Smith 1981). This relationship between global atmospheric circulation and regional oceanographic conditions allows one to predict the position of ancient wind- driven upwelling with paleogeographic reconstructions (Parrish 1982). I will use coastal upwelling to illustrate the physical processes at work behind the phenomenon.

The processes that ultimately drive coastal upwelling are the same ones that drive surface winds, namely the earth's rotation and the differential heating of the earth by the sun. Differential heating sets up a pressure gradient with flow (winds) from the cooler poles to the warmer equator. The effect of earth's rotation (Coriolis effect) modifies the pressure gradient and deflects equatorward-moving air to the left (right) in the Southern

(Northern) Hemisphere. Because of their relatively low heat capacity, landmasses trap or release heat that disrupts zonal atmospheric circulation by creating a land-sea temperature contrast (see reviews in Parrish 1982; Tomczak and Godfrey 1994). Winds formed by 16

the pressure gradient and modified by rotation and land-sea temperature contrast

subsequently drive surface ocean currents.

The transfer of energy from air to water is also influenced by the Coriolis effect,

so that net transport of water occurs perpendicular to the wind direction (Mann and

Lazier 1991). Currents move to the left (right) of the wind in the Southern (Northern)

Hemisphere, and this movement is called Ekman transport. The water must be deeper

than the surface mixed layer for Ekman transport to occur—deep enough for the surface

water to move offshore and for deeper water to move onshore (a bottom Ekman layer)

(Smith 1995). Ekman layers generally are tens of meters deep. Upwelling source water

comes from 100-200 meters depth off of the Oregon and northwest African coasts, but

from only 50-75 meters depth off of the Peruvian coast (Barber and Smith 1981). In

areas where steady winds blow equatorward, parallel to a coast, upwelling can occur

because surface water will move offshore via Ekman transport (Figure 1.2; Mann and

Lazier 1991). During upwelling, a front forms between upwelled (cool) and oceanic

(warm) waters that is distorted by the formation of rings, filaments, and eddies that can

extend productivity hundreds of kilometers beyond the coast (Summerhayes et al. 1995).

Today, coasts along the major eastern boundary currents experience upwelling,

with one exception. The four main areas are found off the coasts of northwest Africa,

southwest Afnca, Peru, and the western United States (Figure 1.2; Mann and Lazier

1991). Australia, however, is a unique case among eastem boundary regions because coastal upwelling does not occur off its west coast. A strong longshore pressure gradient exists in this area, resulting in net poleward transport in the upper layers of water even 17

though winds blow mainly towards the equator. The eastern Indian Ocean connects

directly with the western Pacific Ocean via the Indonesian Islands. Warm water, blown

west by equatorial winds in the Pacific, then travels through the Indonesian Islands and

down Australia's west coast (Smith 1992).

Eastern boundary currents tend to run broad and shallow, resulting in shallower

pycnoclines compared with western boundary currents. A shallower pycnocline means

that nutrient-rich, sub-pycnocline water is more easily upwelled. These characteristics,

combined with upwelling-favorable winds, explain why important upwelling centers are

located on the eastern sides of ocean basins (Marm and Lazier 1991). Wind-driven

upwelling occurs in several other settings as well. Significant coastal upwelling occurs

along the northern coast of Venezuela in the Cariaco Basin (also known as the Cariaco

Trench) (Richards 1975). The Arabian and Somali coasts experience strong seasonal

upwelling during the southwest (summer) monsoon (Barber and Smith 1981), and are the

only east coasts where wind-driven upwelling develops (Mann and Lazier 1991). At the

equator, where the Coriolis effect is zero, the easterly trade winds converge and water is

transported poleward (Gargett 1991), creating a divergence along the equator. This

equatorial upwelling occurs most noticeably in the Pacific ocean, but also happens in the

equatorial Atlantic and Indian oceans (Vinogradov 1981). Upwelling driven by open-

ocean divergence occurs around Antarctica, where the westerlies and polar easterlies

meet (Gargett 1991). In the Gulf of California, wind channels water along the seaway in

what is called channeled-flow upwelling (Parrish 1982). Opposite sides of the Gulf experience seasonal changes in upwelling conditions, following changes in wind 18 direction (Bray and Robles 1993). Physical characteristics of some of the upwelling areas discussed above are found in Table 1.2.

Across the oceans, production is limited by the depth to which light penetrates the water (the photic zone) and by the nutrient supply, which comes mostly from below the photic zone (Mann and Lazier 1991). Where surface waters gain heat (low latitudes), the differential heating creates a density barrier or pycnocline between the warmer waters above and the cooler waters below. While phytoplankton in these regions have ample light for photosynthesis, the inorganic nutrient supply is limited because of the well- defined pycnocline. When surface waters lose heat (usually winter in mid- to high latitudes), surface and deeper waters mix. This provides enough nutrients, but deep mixing can carry phytoplankton below the photic zone where they can no longer photosynthesize (Barber and Smith 1981). At mid- to high latitudes, production is often limited to intense spring blooms when the surface layer warms enough to establish a pycnocline and when nutrients are still plentiful. At low latitudes, production occurs at lower levels year-round because of the stable pycnocline (Mann and Lazier 1991).

Barber and Smith (1981, p. 33) stated that "coastal upwelling is a circulation pattern that overrides both the nutrient limitation of stratified waters and the light limitation of well mixed waters". Vertical movement of water provides abundant nutrients, while advection of surface water helps keep phytoplankton in the photic zone.

Upwelling increases production at the expense of space (Barber and Smith 1981), as these areas are fairly small compared with the rest of the world's ocean area (8.0 x 10^ km^ of upwelling areas compared with 3.6 x 10* km^ of total ocean area, or about 0.2 %; 19

(Pauly and Christensen 1995). However, upwelling increases the frequency and in some

cases the magnitude of productivity events by lengthening the time during which

population growth is favorable. Mid- to high latitudes without upwelling typically have

only one plankton bloom per year, whereas areas with upwelling, such as off the Oregon

coast, can produce several blooms per upwelling season (six months for the Oregon

system). Tropical areas can experience upwelling year round, though with seasonal

variation in strength (e.g., Peru) (Barber and Smith 1981). Organisms respond in

dramatic ways to the increased time available for production, as detailed in the next

section, and this biological response in turn affects the sedimentary and chemical

environments of upwelling areas.

Biological Responses To Upwelling

Nutrients

Nutrient concentration is the primary driver of productivity. In order to stimulate

productivity, upwelled water must contain relatively more nutrients than the displaced

surface water. Generally, elevated productivity occurs if the upwelled water comes from

below the photic zone (below the zone of active nutrient uptake, often below the mixed

layer) as nutrients tend to be depleted at the ocean surface worldwide (Levitus et al. 1993

and references on p. 246). Upwelled, or "new" nutrients come from the upwelling source

water, which in turn usually comes from deep water in the oceans.

The upwelling process itself permits nutrient regeneration by way of a compensatory onshore flow (upwelling water) which acts as a counter-current trap for decomposing particles and regenerated nutrients (Redfield et al. 1963; Barber and Smith 20

1981; Bailey 1991; Toggweiler and Carson 1995). Nutrients in the upwelled water at the

surface stimulate plankton growth. These nutrients are transported out of the offshore-

moving upwelled water as high-density organic matter particles (dead plankton, fecal

pellets). Some of these particles get trapped in the onshore flow as they are moving downward and decomposing. With decomposition, regenerated nutrients are released and upwelled into the surface water again, ready to stimulate new growth (Barber and Smith

1981). Recycling of nutrients keeps production high on the downstream (equatorward) ends of upwelling centers (Bailey 1991).

The Peru and Southwest Africa coastal upwelling systems are an order of magnitude more productive than other coastal systems (Table 1.3). The other coastal upwelling systems have production rates 2-3 times higher or less than non-upwelling systems. In contrast to the Peru and Southwest Africa systems, the other coastal upwelling systems host major river discharges (Alongi 1998), which can decrease productivity by increasing turbidity in the photic zone or by otherwise distiu-bing rapidly growing plankton in upwelling areas. Production in equatorial upwelling zones has been estimated at 176-328 g C m"" y"' (Chavez and Smith 1995), about twice as high as that estimated for other open-ocean areas.

Upwelling does not always result in high productivity. During the 1982-83 El

Nino Event, upwelling-favorable local winds remained constant and even increased off the Peru coast, but productivity decreased because the upwelled water was no longer nutrient-rich. The thermocline and nutricline deepened during this event, cutting off the supply of new nutrients to the surface (Barber 1992). In this case, upwelling occurred. 21

but with nutrient-poor water. However, high nutrient levels do not guarantee elevated

productivity, either. As discussed above, nutrient-rich upwelled water must also remain

in the photic zone so that phytoplankton can thrive. Short-term stratification after an

upwelling event promotes phytoplankton growth and succession. Physical oceanographic

processes—circulation, mixing, turbulence, and water-column stability—greatly affect

productivity levels (Barber and Smith 1981).

Plankton

Phytoplankton convert energy from the sun into carbohydrates with the help of

nutrients. The dominant phytoplankton inhabiting coastal upwelling areas today are

colonial and chain-forming diatoms (Raymont 1980; Rojas de Mendiola 1981;

Summerhayes 1983). These organisms have relatively large cells (5-30 ^m), siliceous

tests, and fast division rates, all of which aid them in utilizing upwelled nutrients quickly.

Siliceous radiolarians and calcareous phytoplankton, including foraminifers and

coccolithophores, are less abundant than diatoms in modem upwelling ecosystems but are

often abundant in sediments (Hutchings et al. 1995).

Most upwelling systems host a particular array of phytoplankton whose

populations are regulated, at least initially, by abiotic factors (Brink et al. 1995). The dominance of picophytoplankton in the equatorial Pacific versus diatoms in the Peru system is at least partly explained by low concentrations of the limiting nutrient, silicate, in the equatorial Pacific (Dugdale and Wilkerson 1998). The Peru system, on the other hand, is not limited by silica so bloom-forming diatoms are plentiful. 22

The zooplankton occupy nanoplankton (<2-20 ^im) through megaplankton (>2000

|im) size classes (Raymont 1983). On the microscopic scale, radiolaria (oceanic) and

tintinnids (neritic, a type of ciliated protozoan) are important prey items for larger

zooplankton. Tintinnids and planktonic foraminifera seem to eat mostly phytoplankton.

Meso- and mega-plankton such as cnidarians and ctenophores can be important predators

of smaller zooplankton. Copepods, euphausiids (mostly oceanic), chaetognaths (all

depths), and protozoa (ciliates, foraminifera, and radiolaria) tend to be the most common

animals found in the zooplankton, with copepods often making up 70% or more of the

fauna. Euphausiids are larger zooplankton (15-20 mm long) and are important in the diets of whales, seals, oceanic birds, and fishes (e.g., herring, sardine. Pacific salmon, tuna). Ostracods can also contribute significantly to the plankton. Planktonic larvae of

benthic and nektonic animals such as molluscs, hydroids, crustaceans, and fishes can make up a large portion of the plankton during blooms (Raymont 1983).

Nekton

An abundant plankton community can support populations of larger, mobile animals. Pelagic and epipelagic shoaling fishes such as sardmes {Sardinops spp.) and anchovies {Engraulis spp.) dominate the nekton in today's upwelling areas (Crawford et al. 1987; Hutchings et al. 1995). These fish are particulate- and non-selective filter- feeders and appear to eat mainly zooplankton, although they can eat phytoplankton when it is especially abundant. Larger pelagic fishes such as horse mackerel compete with shoaling fishes as juveniles and prey on them as adults. Bathypelagic fishes, such hake, also inhabit high-productivity areas (Crawford et al. 1987; Hutchings et al. 1995). 23

Besides fishes, other important higher-level consumers in upwelling systems include squid, pelagic coelenterates, seabirds, and marine mammals (Brodeur and Pearcy

1992). A carbon budget for the southern Benguela system suggested that demersal fishes and squid were important consumers of epipelagic shoaling fishes. Squid are also important consumers in the Benguela region, as well as in the Califomia, Peru, and

Antarctic upwelling systems (Bergh et al. 1985). The Peru, California, and Benguela regions all support large seabird populations (Ainley 1990). These populations can reach huge numbers when prey are abundant. Besides food, a necessary resource for seabird populations is nesting space near the center of food production and isolated from predators. A lack of suitable islands for nesting contributes to the lack of seabirds in the other major upwelling areas (Somali-Arabian and Canary regions) and also limits population sizes (Ainley 1990).

As with seabirds, many cetacean (whale) species flock to high-productivity areas to feed (Slijper 1979; Fordyce 1980; Ray 1981; Gaskin 1982; Reilly and Thayer 1990;

Lalli and Parsons 1993; Reitsch and Veit 1994; Forcada et al. 1996; Jaquet and

Whitehead 1996; Woodley and Gaskin 1996). The mysticetes (baleen whales) seem especially adapted to take advantage of high-density food supplies. Baleen whales usually do not feed unless plankton (usually krill, which are euphausiid crustaceans) densities are high enough to offset the cost of grazing. Baleen whales usually forage when plankton or fish reach densities of 10-500 g/m^ (Dolphin 1987; Piatt and Methven

1992). While odontocetes (toothed whales) are not as directly tied to plankton as baleen 24

whales, the food they eat (fishes, squid) also tends to aggregate in areas of high plankton

concentration (Fordyce 1980; Bloch et al. 1996; Jaquet and Whitehead 1996).

One can expect to find cetaceans (and other marine vertebrates) in coastal and

pelagic regions that experience high productivity either seasonally or year-round. Such

areas include upwelling zones at all latitudes, other areas of divergence (such as

underwater topographic highs), temperate latitudes during the spring plankton bloom (for example, in the North-East Atlantic Ocean), and oceanographic fronts (for example, along western boundary currents or shelf breaks) (Gaskin 1976; 1982; Norris 1983;

Foerster and Thompson 1985; Brown and Winn 1989; Tershy et al. 1991; Ainley 1994;

Olson et al. 1994; Moore et al. 1995). Maps of whale distribution (usually from whaling records) largely reflect major upwelling areas (Townsend 1935; FAO Department of

Fisheries 1972; Gushing 1975; 1982 Figure 90).

As with the plankton (and because of their relationship with the plankton), nekton populations vary widely depending on upwelling intensity. A four-year study of nekton and prey population variability in the Oregon and Washington part of the California upwelling region suggested that changes in a few prey species (euphausiids in this case) affected population sizes farther up the trophic ladder (Brodeur and Pearcy 1992).

Opportunistic feeding habits lend an advantage when prey species fluctuate often, and opportunistic feeders are common in upwelling ecosystems. Oceanographic conditions changed noticeably each year of the study, including a year of strong upwelling followed by a year of El Nino conditions, where upwelled water is warm and nutrient-depleted.

Food web complexity and trophic specialization increased during the El Nifio year 25

(Brodeur and Pearcy 1992). Large-scale environmental changes, such as changes in

regional wind strength (which in turn affects ocean surface temperature, advection, and

turbulence) regulate populations in a large, if indirect, way. Ware and Thompson (1991)

showed that, in general, fish production (measured by sardine and hake biomass) reflects

variation in primary production in the California Current upwelling region.

Large predators such as seabirds and marine mammals are somewhat more

buffered from prey population variability because of their great motility (except for

flightless birds, see Crawford et al. 1987). However, large predators—particularly

opportunistic feeders—tend to aggregate in areas where food is plentiful. These animals

probably stabilize the ecosystem by feeding on multiple trophic levels and by switching

prey taken when prey populations fluctuate (Bergh et al. 1985).

Benthos

The bendios in upwelling systems is also unique. The two main factors

structuring benthic ecosystems in upwelling areas are temperature and oxygen.

Upwelling of cool water allows species typical of lower temperatures to live farther equatorward than in areas without upwelling (Thiel 1978; also true of nektonic and

planktonic species). Consequently, the tropical faunal zone is narrower on the eastern sides than on the western sides of ocean basins (Ekman 1970). For example, warm and cold temperate faunal zones typify the Benguela coast, whereas tropical to subtropical zones are found at corresponding latitudes on the east coast of Afnca (Branch and

Griffiths 1988). Brachiopod faunas differ substantially between the southwest and southeast African coasts (Hiller 1994). The west coast fauna is also depauperate. 26 containing just over half the species of the east coast. At least 3 characteristics of the southwest coast, all resulting from upwelling, contribute to this disparity: cold temperatures (almost 10° C colder on average than the east coast), fine-grained sediments

(most brachiopod species prefer coarse substrates), and high levels of suspended particles. Large populations of the inarticulate brachiopod Discinisca tenuis live off the

Namibian coast. Upwelling systems might be more amenable to settling by inarticulate brachiopods, because these animals can tolerate high concentrations of suspended matter

(Hiller 1994). Benthic suspension feeders can potentially affect nutrient recycling and particulate suspension and sedimentation because of their broad diet and fast grazing rate

(Alongi 1998).

Benthic production generally reflects surface production because of particle transport to the benthic environment (Thiel 1978; Rowe 1985). Benthic organisms in upwelling systems tend to aggregate both nearshore (Amtz et al. 1991) and in areas where organic matter is exported offshore, in contrast to seasonally stratified coastal systems where benthic biomass is highest closer to shore (Rowe 1981; 1985). Benthic standing stock of the continental slope in the Peru upwelling system was higher than in the Gulf of Mexico, a non-upwelling system (Rowe 1985). Both areas showed exponentially decreasing biomass with depth, but differences in absolute biomass reflected the productivity levels. Biomass was about 100 times less on the Gulf of

Mexico (production about 25 g C m'^ y"') slope than on the Peru slope (production 300-

1500 g C m'^ y"') (Rowe 1985). In most upwelling systems, production exceeds even the 27

increased consumption by benthic organisms, so net export of organic matter occurs

(Rowe 1985).

Rowe (1985) recognized two general types of benthic systems in modem

upwelling zones, based on organic matter content and on dominant kinds of metabolism.

The Baja California and Northwest Africa upwelling systems have dominantly

oxygenated benthic habitats. In the Northwest African system turbulence and off-shelf

transport suspend organic particles that support a benthic fauna dominated by filter

feeders (e.g., epifaunal octocorals, feather stars, ascidians, sponges, bivalves, solitary

corals, & polychaetes; also infaunal organisms) (Thiel 1982). In contrast, the Peru and

Benguela upwelling systems contain benthic habitats that are muddy, overloaded with

organic matter, and dominated by dysoxic/anoxic conditions (Rowe 1985). When the flux

of organic matter to the bottom is high, oxygen demand from decay can exceed the supply, leading to oxygen depletion of the sediment or bottom water (Thiel 1978; Tyson and Pearson 1991; Diaz and Rosenberg 1995).

The decomposition of organic matter enhances the oxygen minimum zone

beneath upwelling centers. In upwelling systems overloaded with organic matter, benthic faunas show gradients in diversity and abundance that are related to the degree of oxygenation of the sediment and bottom waters (Mullins et al. 1985; Thompson et al,

1985; Diaz and Rosenberg 1995). The Peru, Southwest Africa, California, and eastern equatorial Pacific upwelling systems all have enhanced oxygen minimum zones (OMZ's)

(Thiel 1978; Diaz and Rosenberg 1995). Benthic macrofaunal diversity tends to be 28

lowest in the core of the OMZ but enhanced at the edges (edge effects) (Mullins et al.

1985; Thompson et al. 1985; Diaz and Rosenberg 1995).

Organisms respond normally to oxygen levels as low as 2 ml/L, but gradually

decrease in diversity and abundance in dysoxic conditions, between 2 ml/L O2 and 0.2

ml/L O2. Even in upwelling areas with high organic matter input to the benthic

environment, very few macroinvertebrates can survive at oxygen levels less than 0.1 ml/L

(0.5 ml/L if in a non-productive area) (Thompson et al. 1985; Diaz and Rosenberg 1995).

The presence of H2S in the sediment further reduces organisms' ability to survive, even at

oxygen levels in the upper dysoxic zone (Diaz and Rosenberg 1995). Studies of modem

organisms indicate that, in general, polychaetes are the taxonomic group most tolerant to

low oxygen conditions, followed by bivalves, and then crustaceans. Macrofauna are

more susceptible to low oxygen than meiofauna or microfauna (Tarazona et al. 1988;

Diaz and Rosenberg 1995).

Low oxygen is responsible for reduced diversity in the OMZ core, while abundant organic matter provides energy and supports organisms that otherwise could not survive

persistent dysoxic conditions (Thompson et al. 1985; Diaz and Rosenberg 1995). Data

from the California, eastern Pacific, and Peru upwelling systems suggest that low-oxygen conditions are persistent both temporally and spatially (Rosenberg et al. 1983; Mullins et al. 1985; Wishner et al. 1990; Hyland et al. 1991; Levin et al. 1991; Diaz and Rosenberg

1995). In upwelling systems, in contrast to other marine areas that experience infrequent dysoxia, the stability of low-oxygen conditions and abundance of organic matter allow low diversity, high abundance benthic communities to survive these harsh conditions 29

(Diaz and Rosenberg 1995). In some areas, fragile mats of bacteria stabilize the environment further. In Peru, mats of Thioploca (purple sulfiir bacteria) dominate low- oxygen zones and concentrate nitrate in the sediment as well as oxidize sulfides (Amtz et al. 1991; Fossing et al. 1995). These mats appear to keep the redox potential discontinuity at the sediment-water interface, preventing the escape of toxic H2S to the water column and enabling some epifauna to live in very low oxygen concentrations

(Savrda and Bottjer 1991; Tyson and Pearson 1991).

While benthic faunas beneath upwelling centers respond to both increased productivity and diminished oxygen supplies, benthic organisms inshore from upwelling centers appear to respond to increased productivity. Only a few studies have concentrated on the nearshore biota near upwelling centers. Inshore from the Benguela system (at tropical-subtropical latitudes), temperate organisms dominate, and kelp beds are common in rocky, sublittoral areas. The fauna is relatively less diverse when compared with subtropical and tropical faunas on Africa's east coast (Branch and

Griffiths 1988). Whereas Benguela intertidal faunas do not change much during El Nino events (Branch and Griffiths 1988), Peru nearshore faunas do, as shown by the dramatic increases in benthic animal abundance and diversity off of Peru during the 1982-83 El

Niiio (Tarazona et al. 1988; Amtz et al. 1991). Secondary benthic production is high in the Peru nearshore system. Whereas some species (such as sulfur-bacteria mats) decline during El Nino years, other species (a species of scallop in 1982-83) thrive. Warmer waters allow subtropical and tropical species to immigrate to the area during El Niiio 30 events. Cooler water faunas return and establish themselves about one year after the event wanes (Amtz et al. 1991).

Biological Responses: Summary

Organisms can affect their environment extensively in upwelling areas, both physically and chemically. During plankton blooms, phytoplankton cell growth can outstrip both grazing and decomposition in the water column, with net sedimentation to the bottom. Fast growth rates and inadequate grazer or predator control overload decomposers who then use up available oxygen, enhancing the oxygen minimum zone and in some cases creating anoxic bottom sediments or waters beneath the production locus.

Some general patterns regarding fauna and flora in upwelling zones are apparent.

Planktonic and benthic organisms in upwelling areas tend to be cooler-water adapted than faunas at the same latitude in non-upwelling areas. Benthic organisms living under upwelling centers tend to be tolerant of low-oxygen conditions. Benthic biomass is related to productivity on the surface, and tends to be high both nearshore and offshore.

Epipelagic shoaling fishes and squid are hallmarks of today's major upwelling systems, and top predators such as seabirds, pinnipeds, cetaceans, and predatory fishes feed in these prey-rich areas. In contrast to seasonal production along coasts, upwelling in low- latitude areas (Peru, Southwest Africa) can operate year-round and generate large quantities of phytoplankton, which can in turn support large populations of cephalopods, fish and other vertebrates. Although these larger animals can migrate significant 31 distances, maps of their distribution coincide with upwelling areas (as well as other, seasonally productive coasts) (Figures 1.3 and 1.4; Gushing 1975).

Little more than general patterns can be drawn from the information available on the biota in upwelling areas, but the patterns can be investigated in more detail in order to shed light on the important controls of upwelling macrofauna. Elevated levels of primary productivity affect not only the biology but also the sedimentology and geochemistry of upwelling zones. The next section discusses how high productivity in upwelling areas creates unique sediments and sedimentation patterns beneath upwelling zones.

Sedimentoloeical Consequences of Upwelling

General Patterns

Upwelling waters are enriched in mineral nutrients such as phosphorus and silica, which promote growth of phytoplankton, ultimately resulting in organic carbon accumulations (Baturin 1983). In areas of persistent upwelling, characteristic sediments are deposited, including combinations of organic-rich shales, phosphorite, chert, and/or glauconite (Baturin 1983; Parrish 1983; 1995; Summerhayes et al. 1995). Trace elements associated with phytoplankton growth or certain oxygen conditions can be enriched in upwelling zones (Baturin 1983; Calvert and Pedersen 1993; Brink et al.

1995).

Studies off of Peru and Southwest Afnca showed that these sediments, where they occur together, exhibit similar facies patterns (Burnett 1980; Calvert and Price 1983). In both cases, organic matter rich in phosphate is concentrated in a central zone, and phosphorite and glauconite pellets are located around the borders of the organic zone. In 32

Peru, the pellets are slightly older than the organic matter (pellets dated by U-series

radiometric methods), and occur along the upper and lower edges of the oxygen-

minimum zone, whereas the organic matter is found in the core of the oxygen minimum

zone (120-385 meters depth; Burnett 1980). Burnett (1980) interpreted these results to

indicate that the zone of maximum upwelling shifted northward during the Quaternary.

This assumes that the pellets were originally deposited as phosphatic organic matter and

became glauconitized under more oxidizing conditions as the zone of maximum

upwelling and the oxygen minimum zone shifted (Burnett 1980).

Glauconitized phosphate pellets also occur off of the Namibian and South African

coasts. The pellets represent reworked intraclasts of phosphatized limestone and provide

a record of past upwelling in the Late Tertiary (Rogers and Bremner 1991). Recent

sediments testify to the continuing high productivity in the area. Sediments in and

around Walvis Bay (inner shelf, northern Benguela region) are enriched in opal (mainly

from diatoms, to >50 weight %), organic carbon (up to 24 weight %), and phosphate

concretions containing abundant fish debris (Rogers and Brenmer 1991). Similar to Peru sediments. Recent phosphate in the Walvis Bay area borders organic-rich diatomaceous ooze (Rogers and Bremner 1991). These observations lead to a predictive model for

upwelling deposits: a regional facies relation of siliceous, organic-rich rock surrounded

by phosphorite (with or without glauconite) strongly indicates high productivity (the Si-

P-C association. Table 1.4; Calvert and Price 1983; Parrish 1983; Parrish et al. 1983). 33

Organic-Rich Sediments

Particulate and dissolved organic matter are produced in large quantities in major

upwelling zones. Much of the production is consumed at the surface and throughout the

water column, but a small fraction (1-25% on the continental shelf) (Legendre and Le

Fevre 1989) rains down to the seafloor, usually in the form of fecal pellets (that often are

ingested and repackaged during descent; Williams et al. 1989). The more organic matter

that gets exported (and the faster), the greater the chances that it will get preserved.

Benguela region sediments contain the highest concentrations of organic matter in the

Atlantic Ocean (up to 24 weight %) and possibly the world (Rogers and Bremner 1991).

Bergh et al. (1985) used a carbon budget model to estimate that perhaps 75% of the

annual primary production in the Benguela system is exported as detritus, either to decay

in the water column or to settle on the bottom. Legendre and Le Fevre (1989) postulated

that the Benguelan food web is less active than those in other upwelling areas (Peru, for

example), so that more production gets exported (Bergh et al. 1985). However,

comparable amounts of organic carbon accumulate on the Peru and Benguelan shelves

(Table 1.4).

The origin of organic-rich marine rocks has been debated for several decades in

the geological literature (Brongersma-Sanders 1948; Demaison and Moore 1980;

Pederson and Calvert 1990; Demaison 1991; Tyson and Pearson 1991; Calvert et al.

1992; Parrish 1995). The discussion centers around what controls the accumulation of organic-rich sediment: preservation by anoxic sediment/bottom waters (insensitive to sedimentation rate) or production (fast sedimentation)? Intense production can create 34

anoxic bottom conditions, complicating the question (see discussion in benthos section).

Another difficulty lies in the different perspectives of researchers working with modem

versus ancient sediments (Tyson 1995).

Supporters of the preservation argument point to the correlation between high

total organic carbon values and low bottom-water oxygen levels as evidence that dysoxic-

anoxic conditions are the primary control on organic matter preservation (Jahnke and

Shimmield 1995; e.g., Tyson 1995). Preservational differences between anoxic and oxic

conditions are most significant at low sedimentation rates, and moderate production and

seasonal stratification are sufficient to produce sediments with enriched organic carbon

(Tyson and Pearson 1991; Tyson 1995). Tyson (1995) noted that in modem

environments, the highest TOC values are associated with high productivity, high

sedimentation rates, and dysoxic-anoxic conditions, confounding the analysis.

Laminated sediments high in total organic carbon (TOC) are sometimes cited as

evidence of the role of preservation in organic-rich sediments. In this view, primary

laminations indicate absence of bioturbating organisms and absence or low levels of

oxygen. However, anoxic sediments accumulate on the Benguela shelf, but they are not

laminated (Summerhayes 1983). Work in the Califomia Borderland Basins and in the

Guaymas Basin showed that macrofaunai organisms can live in very low oxygen

conditions (down to 0.2 mL/L 02; Calvert 1964; Edwards 1985; Mullins et al. 1985).

Edwards (1985) reported the highest TOC values firom sediments slightly disturbed by deposit-feeding polychaetes. In the Guaymas Basin, Calvert (1964) suggested that tube- dwelling worms in the OMZ stabilized laminations in the sediment. Sediment cores from 35

the same localities showed laminations but no worm tubes, indicating low preservation

potential of the tubes (Calvert 1964).

Cuomo and Bartholomew (1991) noted that laminations can result from

compacted fecal pellets of benthic organisms. A coprolitic laminated fabric would

indicate lower dysoxic conditions that support detritus-feeding organisms such as

polychaetes, whereas laminations produced solely from sedimentation with no benthic

activity would contain proportionately more planktonic pellets. Their study indicated that fecal pellets of benthic and planktic origin can be distinguished by inorganic composition

(Cuomo and Bartholomew 1991). Laminated fabrics can also result from wholesale sedimentation of mat-forming diatoms in coastal or open-ocean upwelling zones (Kemp et al. 1996; Pike and Kemp 1999). Laminations must be studied in detail to infer their origin, but overall they are not reliable indicators of benthic oxygen levels.

Proponents of the production argument maintain that high primary productivity

(particularly in upwelling areas) is the main control on sedimentation and, ultimately, preservation of organic matter (Brongersma-Sanders 1948; Summerhayes 1983; Calvert et al. 1992; Parrish 1995). In order to accumulate organic matter beneath upwelling zones, short bursts of high productivity are probably more important than steady production. Massive bursts of phytoplankton overwhelm decomposers in the water column and in the benthos, and allow more of the bloom to accumulate (Summerhayes

1983). The accumulation occurs regardless of oxygen levels at the sediment/water interface. Additionally, large amounts of organic matter, in turn, can create oxygen-poor 36

benthic conditions because of oxygen debt from the decomposers (Summerhayes 1983;

Bailey 1991).

Field evidence supports the importance of primary productivity in supplying

organic carbon to sediments. For example, sediments from areas of different oxygen

levels in the Gulf of California contain comparable amounts of organic carbon (Calvert et

al. 1992). Organic carbon is being deposited on the continental slopes of NW Africa, SW

Africa, and Peru, despite the difference in oxygen levels among these three areas

(Summerhayes 1983). Organic matter is found in surface sediments of the Banda Sea,

which experiences seasonal upwelling, despite lack of a pronounced OMZ (van Waveren

and Visscher 1994). From a literature and database analysis, Parrish (1995) concluded that enhanced productivity could account for over 90% of Phanerozoic oil-prone, organic-rich rocks.

Results of recent studies question the role of anoxia alone in preserving organic carbon. The Black Sea, the largest deep anoxic basin today, contains Holocene-age sapropel overlain by modem coccolith muds (Calvert 1990). The sapropel is sometimes cited as evidence that anoxia plays a dominant role in organic carbon preservation

(Rossignol-Strick et al. 1982). Based on the geochemistry of sediment cores, Calvert

(1990) suggested that the sapropel formed under conditions of increased productivity and oxygen levels as the Black Sea changed from a freshwater to a brackish-water basin.

Because no sapropel is forming today, productivity appears to be the factor triggering organic-rich preservation. 37

Laboratory and field experiments have shown that anoxic conditions do not slow

down decay of organic matter, and that degradation can be just as rapid in anoxic as in

oxic conditions (Heinrichs and Reeburgh 1987; Allison 1988; Alongi 1998). Anoxic

conditions, may, however, promote early diagenesis (Allison 1988) and decrease the

degradation of certain organic compounds (effectively preserving them, Emeis et al.

1991; Tyson 1995). Jahnke and Shinmiield (1995) suggested that the initial stages of

decomposition proceed regardless of oxygen content because the freshest, most labile

organic matter is affected. In this model, oxygen content makes a difference during

remineralization of refractory material at low accumulation rates. Factors besides oxygen

levels or productivity are also involved; controls on fine-grained deposition also affect

organic carbon burial (Jahnke and Shimmield 1995), and the degree of reworking affects

the quality of the organic matter preserved (Emeis et al. 1991).

The production-preservation debate is complex, but regardless, production

ultimately supplies the organic matter and can play a role in its preservation. Today, the

most organic-rich sediments (>20%) are accumulating beneath productive zones in a

variety of oxygen conditions (compare Peru and Benguela). Additionally, very few

totally anoxic basins exist today, and productivity affects their sediments as well (e.g., the

Black Sea, Tyson 1995). These observations argue for an approach that assumes a

primary role for productivity in the origin of organic-carbon-rich sediments.

Biosiliceous Sediments ("Chert")

Summerhayes (1983) noted that diatomaceous ooze is especially characteristic of today's coastal upwelling areas. Diatoms are abundant in the surface waters of all major 38

coastal upwelling areas today, with small, delicate forms most typical of upwelling zones

(Schrader and Sorknes 1991). Diatoms tend to form chains which sink rapidly as the

colony senesces (Raymont 1980). Chain formation contributes to seeding of upwelling

water and to sedimentation of organic matter under upwelling zones, and allows most of

the new production to be sedimented rather than regenerated into nutrients at the surface

(Kiorboe 1993; Hutchings et al. 1995).

Radiolaria (zooplankton) also produce silica in opal form, and their distribution in

modem deep-sea sediments reflects productivity (Leinen et al. 1986). Because opal is

undersaturated in today's oceans, dissolution affects over 95% of the biogenic silica flux

to the ocean floor. Silica that reaches the seafloor generally reflects the distribution of

surface productivity (Herbert et al. 1989; Jahnke and Shimmield 1995). Zooplankton,

particularly larger organisms, package dissolution-prone opal and carbonate particles

(diatom, radiolarian, and foraminiferan tests and coccoliths) into fecal pellets that sink

rapidly through the water column (Porter and Robbins 1981; Pilskaln and Honjo 1987).

Fecal pellets were abundant in sediment traps but absent from core tops at the 3 open-

ocean stations studied by Pilskaln and Honjo (1987), suggesting that pellets remineralize

quickly unless sedimentation (productivity) rates are high.

Phosphorite

Phosphate is a limiting nutrient for organisms. About 95% of phosphate in the

water column is recycled rather than sedimented, and about 5% of phosphate gets

incorporated into marine sediment as organic (fish bones, coprolites) or inorganic

(authigenic, adsorbed, or detrital) phosphate (Follmi 1996). Organic phosphate is labile 39

and breaks down easily (adsorbed inorganic phosphate can be remobilized as well),

whereas inorganic authigenic and detrital phosphate tends to be refractory. Release of

phosphate into pore waters constitutes the basic source of P eruichment in sediments and

appears to be enhanced under anoxic conditions. Conversely, phosphorite precipitation

accelerates under relatively more oxygenated conditions, perhaps explaining why

phosphogenesis seems to occur mainly in the suboxic zone (Follmi 1996). On the Peru shelf, Rowe (1985) observed that phosphate from anoxic pore waters was converting

benthic tests of foraminifera into phosphorite pellets.

Phosphorite is sediment that contains at least 18% P2O5, and that is made up

mostly of organic and authigenic phosphate. Marine phosphorite can occur in the form of nodules, masses, sands, pellets, or oolites (Burnett et al. 1983). Today, phosphorite forms in appreciable quantities in only five regions, four of which are located in coastal upwelling zones (Figure 1.3,2 of the phosphorites forming in coastal upwelling areas are reworked older deposits; only the Peru-Chile and Benguela deposits are of Recent age;

Follmi 1996). In the fifth region, off the eastern coast of Australia, bottom currents rework sediment and concentrate bacterially derived phosphates of Miocene to Recent

Age (O'Brien and Veeh 1983; Follmi 1996). Dispersed, authigenic apatite forms in other areas as well (Mississippi Delta, Long Island Sound), but in quantities well below what one would call a phosphorite (Ruttenberg and Bemer 1993). If the mechanism of phosphate deposition in both upwelling and non-upwelling areas is similar, then formation of phosphorites in upwelling areas is a fimction of the whole sedimentary environment, which concentrates phosphate by a relative lack of terrestrial runoff and winnowing by bottom currents (Ruttenberg and Bemer 1993). Follmi (1996) noted that

current activity is common to all of the major areas of phosphorite formation, with low

net sedimentation rates and high productivity associated with most areas as well.

Phosphorite formation appears restricted to within a few centimeters below the sediment

surface, within the dysoxic zone of organic matter decay. In this case, periodic

winnowing would be important in keeping incipient phosphorites in the dysoxic zone

(Glenn 1990).

Glauconite

While glauconite is not directly related to high productivity, it often forms in association with phosphorite (Birch 1979; Burnett 1980). Glauconite is a greenish, iron-

rich, diagenetic clay mineral (Birch 1979; Monteiro et al. 1983). Glauconite varies in color from pale green to brown, reflecting progressive oxidation (Burnett 1980). It is unclear whether slightly oxidizing or slightly reducing environments favor its formation

(Monteiro et al. 1983). Burnett (1980) reported the highest glauconite concentrations from deeper, more oxic sediments on the Peru slope (0.5 - 1.0 mL/L O2). Mullins (1985) found that the distribution of glauconite corresponded with the upper boundary of the

California OMZ (0.5 mL/L O2). From these observations, it appears that slightly oxidizing conditions favor glauconite formation.

Calcareous Ooze/Chalk

In addition to organic-rich sediment, chert, phosphorite, and glauconite, calcareous sediments can occur in upwelling zones, and seem to be more common in ancient sediments. However, carbonate deposition does not follow the same 41

distributional trends as do the other upwelling facies (Parrish 1983). Today, the main

contributors to carbonate sediments include planktonic foraminifera, coccoliths, and

benthic organisms. Coccolithophores (chalk-producing organisms) become abundant in

less nutrient-rich areas, such as open-ocean upwelling zones (e.g., eastern equatorial

Pacific). Foraminifera are more closely associated with coastal upwelling (Diester-Haass

1978). Coccolithophores can make up a major portion of mature upwelled waters

(stratified, lower-nutrient), as has been observed in tlie Benguela system (Mitchell-Innes

and Winter 1987), but diatoms contribute much more to production and standing crop

(Hutchings et al. 1995). While diatoms contribute abundantly to the sediment during

blooms, foraminifera and coccoliths contribute to the sediment year-round, with possible

significant effects on geologic time scales because of differential preservation (Brink et

al. 1995).

Carbonate deposition in the Benguela system tends to be widespread, with

planktonic foraminifera oozes found seaward of upwelling loci (Bremner 1983; Rogers

and Bremner 1991). Carbonates in the Benguela, however, are not closely associated

with organic carbon distributions, partly because some carbonate is relict, and partly because some of its components (benthic forams) are overwhelmed by contributions firom other sediment producers (diatoms) (Brenmer 1983).

The carbonate compensation depth (CCD) rises where oxygen demand is high

(e.g., high-productivity areas), because decomposer activity increases the amount of CO2 in pore waters (Diester-Haass 1978). Metabolism by sulphate-reducing bacteria, though, can reverse the CO2 increase and even allow carbonate precipitation. In general. 42 carbonates can be preserved if the redox potential discontinuity is above the sediment- water interface (anoxic bottom waters). Less carbonate is preserved if bottom waters are oxic (Diester-Haass 1978). As with any carbonates, upwelling-produced carbonates have a better chance of preservation if they are deposited above the CCD (Krissek and

Scheidegger 1983).

Trace And Minor Element Concentrations

Phytoplankton require certain trace elements for growth, including V, Cr, Fe, Mn,

Zn, Cu, Ni, Mo, and Co (Raymont 1980; Millero 1996). Phytoplankton and other marine organisms concentrate these elements, so the resulting organic matter contains high levels of those elements (Diester-Haass 1978; Piper 1994). Minor and trace metal sediment concentrations are thus highest in areas of very high organic carbon accumulation. Zinc especially has been shown to be concentrated by marine plankton (Calvert and Price

1983). High levels of Ni and Zn were found in organic-rich muds of the Benguela region, but not in organic-rich muds (up to 4 weight %) in the Canary Current region, possibly because of remineralization loss (Diester-Haass 1978). Lithification of phosphate nodules into phosphorite involves recrystallization and organic matter degradation, which lowers the concentration of trace elements in the sediment (Baturin

1983).

Barium is enriched in planktonic foraminifera and in acantharians (a type of radiolarian with a strontium sulphate skeleton). Barite (BaS04) appears to be precipitated

(perhaps biologically mediated) in the presence of decaying siliceous phytoplankton

(Jumars et al. 1989), and this mineral is potentially useful as a qualitative estimate of 43

export production (Brink et al. 1995; Peterson et al. 1995). Barium in sediments of the

eastern equatorial Pacific can be up to 20 times higher than Ba in sediments of adjacent,

less-productive regions (as measured by Barclay) (Diester-Haass 1978). Recent studies

have shown that barite is mostly absent from shelves and in anoxic interstitial waters;

barite is most reliable for deeper waters settings (Von Breymarm et al. 1992).

Uranium enrichment occurs along with phosphorite formation (Diester-Haass

1978) and organic matter accumulation (Baturin 1983). Uranium can be enriched

between 2-200 times the normal levels in sediment (average 6-8 ppm in sediment)

(Baturin 1983). U is a highly soluble anion in oxic conditions, but is reduced to insoluble

uranium oxide under anoxic conditions and in the presence of organic matter (Calvert and

Pedersen 1993).

Framboidal pyrite commonly occurs in upwelling sediments, where it forms

globular clusters and coats and fills organic particles (Baturin 1983). Sulphide

precipitation is especially effective in anoxic waters, where reduced iron species are

abundant (Boulegue and Denis 1983).

Sedimentoloeical Consequences: Summarv

In summary, several different kinds of sediments can be found in upwelling areas,

including organic-rich sediment, chert, phosphorite, glauconite, and carbonate. Organic

matter can contain high levels of bioconcentrated trace elements such as zinc. Today,

only two upwelling systems have the full suite of upwelling sediments (organic-rich sediment, phosphorite, chert, and glauconite). The other systems have some combination 44

of the suite (Table 1.4). Below, I discuss some of the conditions and processes that affect

what sediments are deposited in high-productivity areas.

Sources Of Variable Sedimentation

Variability is a common theme in upwelling sediments (Table 1.4; Summerhayes

et al. 1995). Comparison and contrast of the upwelling areas listed in Table 1.4 can aid in

identifying the factors that contribute to depositional variability.

Of the four main coastal upwelling zones, only the Benguela and the Peru-Chile

areas contain the full suite of upwelling-associated lithologies (organic-rich sediment,

phosphorite, chert, and glauconite). These areas are both situated at tropical-subtropical

latitudes and are offshore of arid regions that receive some eolian sediment but relatively

little runoff (reflected in low Al contents compared with non-upwelling areas for

Namibia, Calvert and Price 1983). These are also the upwelling areas with the highest

TOC contents. The coastal upwelling areas located in temperate humid or tropical seasonally wet climates (California, Northern Arabian Sea) tend to have the lowest organic carbon contents and also do not contain phosphorite. Thus, climate is one of the main factors that shapes the sediment record of an upwelling zone. Biogenic sediments of an upwelling zone in an arid region are less likely to be diluted by terrestrial runoff.

Phosphorite formation is also favored by high productivity but low sedimentation rates

(Follmi 1996), such as might be found around the edges of an upwelling zone in an arid region. On the other hand, runoff, particularly if rich in dissolved nutrients, can enhance productivity if the water column is adequately stable and transparent (though runoff is likely to be nutrient-poor in tropical areas, Raymont 1980). Runoff raises the 45

sedimentation rate, which can help preserve biogenic sediments (van Waveren and

Visscher 1994), but those sediments are inevitably diluted, so biogenic sediment

concentrations will be less in humid than in arid regions. Also, in more humid areas such

as the California system, organic-rich sediment is hemipelagic or concentrated farther

from shore, whereas abundant organic matter is found near the coasts in the Benguela

and Peru-Chile upwelling systems (Summerhayes 1983).

Climate not only affects upwelling sediments in terms of dilution of biogenic

sediments, but also in terms of upwelling seasonality and strength. Year-round upwelling

provides more opportunity for favorable planktonic bloom conditions and higher

production. This is reflected in the abundance and diversity of biogenic sediments in the

Benguela and Peru-Chile zones, both of which experience year-round upwelling (Tables

1.2 and 1.4). Seasonal upwelling, on the other hand, provides less time for blooms to

develop and generally results in lower production, as shown by the estimates of primary

productivity and TOC contents for the seasonally productive California. Cariaco Basin,

and Somali coastal upwelling zones (Tables 1.2 and 1.3).

While climate is a major influence on upwelling sedimentation, it is not the only

one. The Northwest African upwelling zone is located offshore of an arid region and experiences year-round upwelling, but few upwelling-related sediments are deposited there compared with the Benguela and Peru-Chile areas. Organic carbon deposition in the northwest Afncan system is concentrated on the lower slope rather than the shelf and

upper slope as in other upwelling systems (Thiel 1982). The Northwest Afncan shelf is quite shallow across its entire width (shelf break at 105 m, while world average is 132 46 m), leaving it susceptible to swells and bottom currents that can winnow out fine-grained material and deposit it in deeper waters (Thiel 1982; Summerhayes 1983). Waters over this shelf are turbulent because winds are strong but variable. Turbulence reduces productivity because phytoplankton spend less time in the photic zone (deep mixed layer;

Summerhayes 1983; Bailey 1991); a deeper mixed layer can also enhance organic matter degradation (Tyson and Pearson 1991). In this case, shelf depth is an important factor that affects the sediment record of Canary Current upwelling and causes the sediments to differ from other upwelling regions in dry, tropical climate zones.

The preceding comparison of coastal upwelling zones shows that climate and shelf depth are majorfactors affecting variability of sedimentation in upwelling areas.

Aridity and seasonality are the aspects of climate that most affect the sediment signal.

Upwelling areas located offshore of arid regions are more likely to have relatively undiluted biogenic sediments. Phosphorite seems to form only in these regions, where productivity is high but sedimentation rates around the edges of the upwelling area are low. Seasonality affects the upwelling signal by determining the time available for production and the production intensity. Year-round upwelling provides more time for biogenic sediments to be produced and deposited. Shelf depth modifies the climate influence in that a shallow shelf can result in turbulence, which reduces productivity levels and winnows fine-grained biogenic sediment off the shelf to the lower slope.

Understanding the factors that lead to different sedimentation patterns in upwelling zones aids in developing criteria for recognizing ancient upwelling systems.

By analogy with the distribution of modem upwelling sediments, the full suite of 47 upwelling-associated sediments (organic-rich sediment, phosphorite, chert, and glauconite) is expected only in regions where runoff is minimal and upwelling is year- round (to allow for maximal biogenic sedimentation)—arid, tropical regions. Upwelling today occurs in a variety of climate zones, so additional criteria are needed to help identify upwelling in other areas. 48

ANCIENT UPWELLING SYSTEMS

By studying the biology, sedimentology, oceanography, and geochemistry of

modem upweiling systems, we can predict what aspects of upweiling might be preserved

in the rock record. It must be emphasized that no single definitive criterion for upweiling

exists; multiple indicators are necessary to establish upweiling for an ancient deposit

(e.g., Calvert and Price 1983; Parrish and Gautier 1993; Peterson et al. 1995;

Summerhayes et al. 1995). The following discussion focuses on aspects of sediments and

biota observed in modem upweiling systems that could be preserved over long periods of

geological time (millions of years).

Preservable Aspects of Upweiling Sediments and Biota

Table 1.5 lists the upweiling conditions and preservable sedimentary and

macrofaunal characteristics of four modem upweiling zones. These upweiling zones

represent a continuum of upweiling intensity.

Upweiling deposits have characteristic geometries that could be recognized in a

regional facies analysis. Coastal upweiling deposits are generally parallel with the

shoreline. The concentric arrangement of facies in the Si-P-C association, as found in the

Pern and Benguela systems, is another preservable characteristic. In the geologic record,

one could expect to find a belt of organic-rich rock surrounded by phosphatic and

glauconitic sediments (biogenic silica or chalk could be interspersed throughout the other

facies).

Preservable lithologic characteristics include the Si-P-C association or single occurrences of organic-rich rocks, biogenic silica, or phosphorite (glauconite in isolation 49

does not necessarily indicate high productivity). Three of the four upwelling zones listed

in Table 1.5 have sediments with TOC contents above the world average for shales (1

weight %; Boggs 1992). Along with organic-rich rocks, one would expect higher levels

of bioconcentrated trace metals such as zinc (Piper 1994). Organic-rich fecal pellets are

abundant in many modem upwelling areas (Benguela, Peru, and California systems of the

areas listed in Table 1.5). Fecal pellets may be important components of many ancient

organic-rich sediments (Porter and Robbins 1981).

In areas where the full suite of upwelling lithologies is absent, oxygen-minimum-

zone edge effects are potentially useful indicators of high productivity, as demonstrated

for the California upwelling zone (Thompson et al. 1985; Vercoutere et al. 1987). Edge effects are manifest in the central California OMZ by downslope changes in sediment grain size and type, fecal pellet abundance, and benthic diversity and abundance. The

upper edge of the OMZ is characterized by coarse, glauconitic sediment and abundant asteroids and ophiuroids. The core of the OMZ contains bioturbated, fine-grained sediments relatively depleted in TOC; it also contains a diverse but low-abundance benthic foraminifera assemblage and abundant echinoids and hermit crabs. The lower edge of the OMZ contains bioturbated sediments with high TOC, abundant fecal pellets, and abundant asteroids and ophiuroids (Thompson et al. 1985; Vercoutere et al. 1987).

Polychaete abundance peaks at the upper OMZ edge, while echinoderm abundance peaks at the lower OMZ edge; these peaks correspond to peaks in TOC contents in OMZ sediments (Thompson et al. 1985). While polychaetes are less likely to be preserved in the geologic record, echinoderms such as asteroids could be preserved. 50

Because upwelling typically involves cool water, a preservable biological aspect of upwelling is the presence of cool-water organisms in regions otherwise interpreted as subtropical or tropical. Cool-water phytoplankton, zooplankton, and benthic organisms

(e.g., ostracods and brachiopods in the Bengeula system, Hiller 1994; Dingle et al. 1996) are typical of upwelling systems today.

Food is readily available in upwelling areas and is reflected in the ecosystem structure. Opportunistic species are abundant (California system, Brodeur and Pearcy

1992), as are large pelagic predators (whales feed in all major upwelling areas today).

Today, filter- and particle-feeding fish are dominant components of large upwelling systems (e.g., sardines or anchoveta in the Peru and Benguela systems). Non-symbiotic suspension feeders, such as brachiopods or bryozoans, also tend to be characteristic of high-productivity environments (Brasier 1995). An abundance of these kinds of taxa in the fossil record could indicate high productivity conditions.

Mass kills of fishes associated with blooms of dinoflagellates occur in upwelling centers of the Benguela system (Brongersma-Sanders 1948); such mass kills could be preserved (in the case of the Benguela mass mortality, fish remains would be associated with diatomaceous, organic-rich sediments). Brongersma-Sanders (1948) suggested that upwelling environments could have been important sites for mass kills over geologic history.

Evaluation

A comparison of the preservable features of the four upwelling zones in Table 1.5 shows that the Peru, Benguela, and California upwelling systems would be the easiest to 51

identify as upwelling zones if a geologist in the future was studying their sedimentary and

fossil records (assuming that all of the features were preserved). Each of these three

regions contains a variety of biological and sedimentological upwelling features. The

California system lacks the Si-C-P association but has enough other features to make the

upwelling identification, including OMZ edge faunas and sediment distributions,

moderately high TOC, biogenic silica (though much diluted compared to the Peru and

Benguela systems), cool-water organisms, abundant pelagic taxa, non-symbiotic

suspension feeders, and abundant large predators. The Northwest African upwelling

zone would be the most difficult to identify, especially if no fossils were available.

Northwest Afncan upwelling is less productive than the other upwelling zones, and the

OMZ is weakly developed. In this case, biotic features of the system, such as the

presence of cool-water taxa, abundant pelagic and planktic taxa, and the presence of non-

symbiotic suspension feeders would be helpful in identifying the area as an upwelling

zone.

The variable nature of upwelling sedimentation means that less-intense upwelling

zones are more difficult to identify based on their sediments alone, as the Northwest

Afnca example above showed. Less-intense or small-scale coastal upwelling usually is

less widespread and thus has less chance of getting preserved. Additionally, less-intense

upwelling produces fewer biogenic sediments. For example, sediments produced in the

Washington-Oregon upwelling area (northern part of the California system) do not differ

much from those produced off of a stratified shelf system such as the New York Bight.

Primary productivity and organic carbon contents are similar in both areas (200-300 52

gC/m^; <0.5 -1% Corgon the shelf; 1-3% on the slope). The New York shelf contains

terrestrial as well as marine organic matter, and the Columbia River in the Washington

area contributes considerable sediment and terrestrial organic matter to the shelf (Walsh

1981; Carpenter and Peterson 1989; Landry and Hickey 1989). Off Oregon, the OMZ is

quite deep (700-1000 m) and oxygen levels are higher than in the California part of the

zone (though still down to 0.3 mL/L O2) (Krissek and Scheidegger 1983). Perhaps the

only differences that could be detected in the geological record would be OMZ edge

effects in the Oregon region (edge effects have been documented in the central California

part of the same upwelling system), and regional facies differences in the locus of

organic-rich deposition (estuarine/slope off New York, slope off Oregon). Both regions

are influenced by cool-water currents (NY—Labrador Current, OR—California Current),

so it is unlikely that temperature-based faunal differences would exist.

The Oregon-New York comparison described above suggests some minimum

requirements for detecting ancient upwelling. Sediments that contain only slightly

enriched TOC could result from production associated with seasonally stratified shelves

or from upwelling. More specific information is needed on faunal composition and ecology in order draw more specific conclusions. On the other hand, the presence of the full suite of upwelling sediments (organic-rich sediment, phosphorite, chert, and glauconite) seems to occur exclusively ui upwelling areas; today this facies association is known only from the Peru and Benguela upwelling systems (Table 1.5). Preservable macrofaunal indicators of upwelling, described above, are necessary in order to identify upwelling in sediments that fall between the Oregon and Peru-Benguela endmembers. 53

Note also that all of these macrofaunal indicators are present in the Peru and Benguela

upwelling systems as well.

Examples of Ancient Upwelling Systems

As most work on upwelling indicators has focused on sedimentological and

microfaunal upwelling indicators (Table 1.1), it is not surprising that several of the most

well-known ancient upwelling areas were identified based on the presence of the Si-C-P

facies association. Three of these well-known deposits include the Miocene Monterey

Formation of California, the Triassic Shublik Formation of Alaska, and the Permian

Phosphoria Formation of the western Rocky Mountain region (Table 1.6).

The Miocene Monterey Fonnation outcrops along much of the California coast and contains high-productivity facies including siliceous, phosphatic, and calcareous mudstones, and carbonaceous marls (see papers in Garrison et al. 1981; Isaacs 1987;

MacKinnon 1989b; 1989c). These sediments were deposited extensively in pull-apart basins that formed along the California coast in the late Oligocene - late Miocene, as the

East Pacific Rise collided with the North American Plate (Pisciotto and Garrison 1981).

Terrigenous runoff was low at this time and hemipelagic sedimentation dominated. Piper and Isaacs (1995) determined that minor elements (Cd, Cu, Mo, Se, Zn, Fe) in the marine fraction of the rocks had interelement ratios similar to those in modem plankton, consistent with a biogenic origin.

In addition to the Si-P-C association, the Monterey Formation also contains abundant diatoms, calcareous microfossils (coccoliths and foraminifers) and fish remains.

The calcareous microfossils are most abundant in the lower part of the section and 54 probably resulted from a less nutrient-rich but stable water column (Pisciotto and

Garrison 1981). Abundant marine vertebrate remains (whales and pinnipeds) are present

(Barnes 1977) as well as possible fossilized bacterial mats, which might have enhanced phosphatization (Reimers et al. 1990) and provided a substantial portion of the oil- forming organic matter in this petroleum-rich formation (TOC varies from 0.5-34 weight

%) (Williams and Reimers 1983; Isaacs 1987). The siliceous fades alternates between laminated and homogeneous, bioturbated intervals; massive and poorly laminated beds contain more organic matter than do well-developed laminated beds (Isaacs 1983).

Overall, the Monterey Formation records moderate-high, upwelling-induced productivity which increased in the middle Miocene (siliceous and phosphatic fades) and then decreased in the Pliocene as sea level fell and terrigenous input increased (Table 1.6;

Pisciotto and Garrison 1981; Barron 1986).

The Shublik Formation comprises sediments that were deposited along a south- facing passive margin on the North Slope of Alaska during the Triassic (Parrish 1987a;

Kupecz 1995). Though heterogeneous (Parrish 1987a), the distinctive nature of the rocks and fossils in this unit, plus its widespread occurrence across northern Alaska (Detterman et al. 1975), readily allow fades mapping. From north to south, lithofacies include non- glauconitic sandstone, glauconitic sandstone and siltstone, phosphate-nodule-containing siltstone and silty limestone, and organic-rich, black limestone, marl, and mudstone

(Parrish 1987a). Contemporaneous siliceous rocks of the Otuk Fm. are located farther south in the Brooks Range (Mull et al. 1982). Temporally, the Shublik Fm. comprises 3 55

informal members: siltstone, limestone and dolomite, and clay shale (ascending order)

(Detterman et al. 1975).

Fossils are abundant in the phosphatic and organic-rich facies of the Shublik Fm.

(Detterman et al. 1975; Parrish 1987a). Bivalves, gastropods, ammonoids, brachiopods,

and ichthyosaur bones are common in limestones of the phosphatic facies (Detterman et

al. 1975; Parrish 1987a). In the organic-rich facies, ichthyosaur bones are again

common, and the invertebrate fauna, while severely reduced in diversity, contains very

abundant bivalves solely of the genera Halobia and Monotis. These bivalves have thin,

flat shells that comprise the fine laminations of this facies. Impressions often cover

bedding planes, and Parrish (1987a) interpreted the bivalve abundance as indicating high

productivity.

Permian rocks from the west-central U.S. (UT, WY, MT, ID) are so phosphatic

(up to 35 weight %, Piper and Medrano 1994) that they were named the Phosphoria

Formation. This formation and its equivalents outcrop extensively in the region

(McKelvey et al. 1959; Wardlaw and Collinson 1986; Sheldon 1989). From east to west, lithofacies include red beds (Goose Egg Fm.), carbonates (Park City Fm.), and phosphatic shale, siltstone and chert (Phosphoria Fm.) (Wardlaw and Collinson 1984). These rocks were deposited at equatorial paleolatitudes in a foreland basin between the North

American passive margin and the Antler orogenic belt (Wardlaw and Collinson 1986).

Using conodont and brachiopod assemblages, Wardlaw and Collinson (1984) showed that the phosphatic facies generally corresponds to cool and nutrient-rich offshore shelf environments. Arctic brachiopods, such as Neospirifer striatoparadoxus. 56

dominate these deeper carbonate ramp facies, while fusulinid foraminifers, thought to

live in warm waters, are absent from Phosphoria rocks (Wardlaw and Collinson 1986).

McKelvey et al. (1959) noted that the phosphorites often contain nothing but inarticulate

brachiopods and some fish debris. Acritarch remains (phytoplankton; Jacobson et al.

1982) and cephalopods (including squid; Yochelson 1963) are also found in the

phosphorites. Corals are rare, and molluscs occur most abundantly in carbonate facies.

Pectinid clams are distributed in patches across the carbonate bank, whereas cephalopods

are found in the offshore facies. Bryozoans are abundant in carbonates, and crinoid

columnals, siliceous sponge spicules, burrows, and fish debris are common (Wardlaw and

Collinson 1986).

The abundance of phosphorite and organic-rich sediment, presence of cool-water species (and absence of warm-water species), and abundance of brachiopods, molluscs, and fish suggest high-productivity in an upwelling zone. Epifaunal and pelagic organisms dominate the assemblages (Yochelson 1963). The abundance of inarticulate

brachiopods is reminiscent of the brachiopod fauna inhabiting the Benguela upwelling

region today (Hiller 1994). Offshore assemblages in phosphatic and shaly sediments differ from those in carbonate sediments (Yochelson 1963), perhaps because of cooler temperatures and influence firom the OMZ (Wardlaw and Collinson 1986).

All three of these ancient upwelling deposits contain economically important deposits of either petroleum (Monterey, Shublik, Phosphoria), phosphorite (Phosphoria), diatomite (Monterey), or trace metals (V in Phosphoria) (Pisciotto and Garrison 1981;

Parrish 1987a; Sheldon 1989; Piper and Medrano 1994). Just as importantly, each 57

contains abundant and distinctive fossil assemblages that can be related to high surface

and benthic productivity and low-oxygen conditions in the benthic environment. Each of

these formations contains depauperate benthic faunas in outer shelf, slope, or basinal

environments. Epifaunal benthic organisms tend to predominate (e.g., brachiopods in the

Phosphoria, Yochelson 1963), and pelagic and/or planktic organisms are common. Cold-

water organisms support upwelling of deeper, colder water (e.g., brachiopods in the

Phosphoria, Wardlawand Collinson 1986).

In each of the preceding examples of ancient upwelling deposits, sedimentologic

and biologic indicators of upwelling are numerous. When sedimentologic indicators of

upwelling are few or lacking, however, macrofaunal features alone can be useful in

assessing the role of upwelling. Analysis of biologic indicators suggests that Pliocene-

age sediments in southem Florida represent the effects of upwelling in shallow-marine sediments (Allmon 1993; Allmon et al. 1996). Of the 5 Pliocene units, only the oldest,

the Upper Bone Valley Formation, contains abundant phosphate in the form of sands and

gravels (Allmon et al. 1996). The Bone Valley Formation is thought to represent

nearshore and estuarine environments (Cathcart 1989). Lithologies in units above the

Upper Bone Valley Formation include sands and limestones with abundant, dense fossil assemblages (Allmon 1993).

Many of the organisms found in the Pliocene beds no longer inhabit Florida waters or are much less common, including many kinds of cetaceans, pinnipeds, seabirds

(especially cool-water species), and molluscs (notably turritellid gastropods, which today inhabit cool and/or nutrient-rich waters) (Allmon et al. 1996). One bed in the sequence 58 contains a monospecific assemblage of thousands of bones of an extinct cormorant whose relatives currently live only in the northeastern Pacific Ocean (Emsiie 1995). Calcareous algae, which typically live in warm waters, are absent from these sediments, and the dominant corals do not form reefs but rather small, isolated colonies which might have lived in turbid, cool waters (Allmon et al. 1996). Limited isotopic evidence from bivalves and gastropods suggests seasonal upwelling events, with enrichments in 5'^0

(cooler water) corresponding to depletions in S'^C (nutrient-rich; Jones and Allmon

1995). Abundance and diversity of the marine vertebrates and cooler/nutrient-rich invertebrate taxa decreased after the Pliocene, when upwelling along western Florida declined with the closing of the Central American Seaway to form the Isthmus of Panama

(Allmon etal. 1996). 59

SUMMARY

Biological indicators provide key evidence for oxygen-minimum zones,

upwelling of cool water, and iiigh productivity, as illustrated by fossils in the Phosphoria

Fm. and the Florida Pliocene (Wardlaw and Collinson 1986; Allmon et al. 1996).

Ancient upwelling environments can be identified, provided that enough biological, sedimentological, and geochemical evidence is preserved to make the determination (note indicator distributions in Table 1.5). This is especially important when dealing with past systems, which sometimes leave a record suggesting very different environments from what we observe today. Parrish (1982) noted that while little organic carbon accumulates today beneath open-ocean divergences, this type of upwelling could have been important in the past over shallower, epicontinental seas.

In this chapter, I have described how upwelling systems work and their signature in the geologic record. I emphasized the role that organisms play in structuring the upwelling environment, and presented criteria that can be used to identify ancient upwelling areas. In Chapter 2,1 apply these ideas to the Upper Triassic Luning

Formation, a rock unit with abundant ichthyosaurs that presumably fed within the Luning depositional basin. 60

Upwelling front North Warm

Cool

Coast

Shelf-edge upwelling East Ekman transport

Shelf

Deep IJ compensatory current

FIGURE 1.1. Simplified view of a coastal upwelling system in the Northern Hemisphere. Modified from Mann and Lazier, 1991 with additional information from Summerhayes et al., 1995. California- Northwest Northern Oregon Africa I Arabian V Sea summer^^l Cariaco K^Winter Equatorial Pacific-y///7/y///// - Summer

Peru Bcnguela

Anlarctic divergence

FIGURE 1,2 Today's major upwelling areas. Arrows indicate prevailing winds. Coastal upwelling areas are stippled; open-ocean upwelling is hatched. After Mann & Lazier, 1991, with additional information from Bray & Robles, 1993; Gargett, 1991; Parrish, 1982; and Richards, 1975. e,

60°

30°

0° h 30° 1

60°

120° 180° 120° 60° 0° 60°

FIGURE 1.3. Global distribution of present-day phosphorite formation (stars), relict phosphorites on the seaftoor (triangles), coastal upwelling (bold lines), and guano island distribution on continental shelves (filled circles). Phosphorites forming from Recent biogenic material are found only in the Peru-Chile and Benguela upwelling systems. Base map and guano distribution from Gushing, 1975; phosphorite localities from Follmi, 1996; coastal upwelling localities from Follmi, 1996 and Mann and Lazier, 1991. The Gulf of California and Cariaco Basin upwellings are not shown (see Figure 1.2). S 63

TABLE 1.1. Representative references on different aspects of upwelling. Much more work has been done on the sedimentology, geochemistry, and micropaleontology of ancient upwelling zones than on the macropaleontology. While this table does not include every available reference to ancient upwelling, it is representative of the proportion that focus on the different aspects of upwelling.

Focus of article Representative references Macropaleontology Fordyce, 1980; Farrish and Parrish, 1983; Fordyce, 1991; Allmon, 1993; Allmon et al., 1996

Micropaleontology Diester-Haass and Schrader, 1979; papers in Suess and Thiede, 1983; papers in Thiede and Suess, 1983; Ingle, 1987; Diester- Haass et al., 1988; Altenbach and Samthein, 1989; papers in Summerhayes et al., 1992; Eshet and Almogi-Labin, 1996

Sedimentology and Samthein et al., 1981; Einsele and Wiedmann, 1982; Parrish, Geochemistry 1982; Parrish and Curtis, 1982; Einsele and Wiedmarm, 1983; Lindstrom and Vortisch, 1983; Parrish et al., 1983; papers in Suess and Thiede, 1983; Summerhayes, 1983; papers in Thiede and Suess, 1983; Gardner et al., 1984; Riggs, 1984; Glerm and Arthur, 1985; Lagoe, 1985; Parrish, 1987b; Parrish, 1987a; Follmi, 1989; Gleim and Arthur, 1990; Dymond et al., 1992; Lyle et al., 1992; papers in Sunmierhayes et al., 1992; Almogi-Labin et al., 1993; Parrish and Gautier, 1993; Dean et al., 1994; Glerm et al., 1994a; Glerm et al., 1994b; Schoell et al., 1994; Parrish, 1995 TABLE 1.2. Physical oceanographic characteristics of major upwelling areas in today's oceans.

Typical Sea Surface Primary Temperature During Productivity Upwelling area Location Upwelling duration Upwelling Periods^ gCm-^f' year-round, varying southwest African coast, 18- intensity (northern part); 7-10° C(LUderitz area) 1000-2000 Benguela Current 35° S lat. seasonal (southern part)

central North American coast, seasonal, varying 7-10°C (central California Current 150-200 30-45° N lat. intensity* California coast area)

northwest African coast, 10- year-round, varying 15-I7°C (near Cap Canary Current 200-500 40° N lat. intensity Blanc)

22-23° C (over 28° northern Venezuela coast, 11° seasonal, late winter during non-upwelling 90-250 Cariaco Basin NIat. periods)

Pacific Ocean, 8-12° N, 6-8° Eastern Equatorial year-round, varying 22° C (eastern tropical S, 90-130° W (open -ocean 175-300 intensity Pacific) Pacific divegence)

west South American coasts, year-round, varying Peru Current 15.5-17° C (near 15° S) 1000-2000 0-20° S intensity

east African & southeast Arabian coasts (summer); seasonal—varies with 19-22° C (near Oman) 175-200 Somali Current west Indian coast (winter); the monsoon I0-20°N •Winds change over short time periods (days) off the Oregon coast, but are steadier off the California coast (Brink et a!., 1995) tAverage sea surface temperatures: >28° C in equatorial regions, 0° C in polar regions (Tomczak and Godfrey, 1994, figure 2.5). Sources; (Anderson et al., 1992; Barber and Smith, 1981; Brink et al., 1995; Chavez and Smith, 1995; Mann and Lazier, 1991; Minasetal., 1982; Moody etal., 1981; Richards, 1975; Shannon, 1985; Smith, 1995) 65

TABLE 1.3. Primary productivity estimates for representative areas of the world's continental shelves (modified from Alongi, 1998; tChavez and Smith, 1995; •Richards, 1975). Primary Latitude Major Production (°) Region River (g C m"^ yr"')

Meridional Coastal Upwelling (Eastern Boundary Current) 0-30 Ecuador-Chile 1000-2000 Southwest A frica 1000-2000 Northwest Africa 200-500 Baja California 600 Somali coast Juba 175 Arabian Sea Indus 200 30-60 California-Washington Columbia 150-200 Portugal-Morocco Tagus 60-290

Meridional Dynamic Upwelling (Western Boundary Current) 0-30 Brazil Amazon 90 Java/Banda Seas Brantas 110 South China Sea Mekong 215-317 West Florida shelf Appalachicola 30

Zonal Coastal Upwelling 0-30 Cariaco Basin Manzanares 90-250*

Equatorial Open-Ocean Upwelling 0-5 Eastern Equatorial Pacific 175-300^

Mesotrophic Systems 30-60 Australian Bight Murray 50-70 Southern Chile Valdivia 90 Southern Mediterranean Nile 30-45 Bering Sea Kuskokwim 170

Phototrophic Systems 60-90 Beaufort Sea Mackenzie 10-20 Chukchi Sea Yukon 40-180 Weddell-Ross Seas 12-86

Eutrophic Systems 30-60 Mid-Atlantic Bight Hudson 300-380 Baltic Sea Vistula 75-150 East China Sea Yangtze 170 Louisiana/Texas shelf Mississippi 100 TABLE 1.4. Geologic features of modem upwelling areas Upwelling area Geographic features Locus of organic Sediment types TOC Sedimentary structures Additional deposition deposited weight % comments Benguela region Passive, shelf & slope. Shelf, near the coast; marine organic 1-15%; up fecal pellets, inner shelf platform in Slope matter (OM), opal, to 24% phosphate/glauconite some areas; shelf 100- PiOj, glauconite. pellets & nodules 160 km wide carbonate California north—narrow shelf. Slope, basins marine OM, 2-4.5% fecal pellets, trace TOC <0.5% Current subduction zone glauconite. fossils, laminations off Oregon carbonate, turbidites (below 0.2 mL/L Oi), coast south—pull-apart locally (contain result from seasonal basins; continental terrestrial OM) terrestrial runoff (STR), borderland which dilutes OM Cariaco Basin Pull-apart basin, silled. Basin marine and 2.5-4% mm-scale laminations Anoxic waters with 2 sub-basins 1400 terrestrial OM (from STR), fecal below about m deep, 900 m ridge pellets, microbioturb- 300 m depth separates sub-basins ation, microturbidites Eastern Deep ocean basin; Basin, seamount diatom mats & thin 3-3.5% Laminations (diatom mat sed. rates Equatorial oxic benthic slopes that intersect layers of mixed (one mats), trace fossils; >160 cm/ky Pacific conditions the OMZ calcareous & sample); mats result from frontal (lOx higher siliceous <0.25- zone convergence, not tlian normal microfossils 0.5% upwelling per se upwelling) Northwest Africa Passive, shelf & slope; Slope between 1000- carbonates, marine 1-4% coarse material high turb­ shelf shallow (50-100 2000 m, small OM, some eolian predominates on the ulence and m) & wide; shelf edge nearshore basins dust shelf turbidity shallow (105 m) Peru-Chile Active, shelf & slope. Shelf-slope marine OM, opal. 1-10%; up laminations, trace narrow shelf PiOj, glauconite. to 22% fossils, phosphate/glau­ carbonate conite pellets & nodules Northern Arabian Active & passive. Shelf-slope marine OM, carbon­ 1-4 %; up laminations Opal at depth; Sea shelf & slope ate, clastics, opal to 11% diluted & (W India) dissolved higher up? References: Berger and Herguera, 1992; Calvert and Price, 1983; Diester-Haass, 1978; Emeis et al, 1991; FUtterer, 1983; Gorsline et al., 1996; Hughen etal., 1996; Kemp, 1995; Kempetal., 1996; Levin etal., 1991; Mullins et al., 1985; Richards, 1975; Rogers and Bremner, 1991; Schulzet al., 1996; Seibold, 1982; Summerhayes, 1983; Thompson etal., 1985. 67

TABLE 1.5. Upwelling conditions and preservable biologic and sedimentary upwelling indicators in 4 Recent, major upwelling zones. Peru, Benguela California Northwest Afnca Most intense Less intense Less intense Upwelling conditions Year-round coastal upwelling X Seasonal coastal upwelling Minimal runoff X Low-moderate runoff High seasonal runoff X Well-developed OMZ X X Less well-developed OMZ Weak bottom currents X Moderate-strong bottom currents Strong bottom currents

Indicators expected in the geologic record Cool-water organisms X X X Abundant planktic ScJor pelagic fossils X X X Opportunistic species X X 9 Non-symbiotic suspension feeders + lack of reefs X X X Large, pelagic predators X X 7 Depauperate benthos X X Mass kills X X X Deposits aligned parallel with coast X X X Si-P-C facies geometry X Biogenic silica X Phosphorite X Biogenic carbonate X X X Organic matter enrichment X X X Glauconite X X High levels of trace elements X X 7 OMZ edge effects X X 9 TABLE 1.6. Comparison or4 ancient upwelling deposits'

DEPOSIT FACIES" MICROI-ADNA MACROFAUNA GEOCHEMISTRY Monterey Fm. Miocene, 17.5-6 Siliceous" Phytoplankton: Pelagic: TOC: Ma'", CA coast Phosphatic marlstone Diatoms Fish & marine mammals— 5-34 wt.% in phosphatic 300-3000 m thick cocco1 ith-foram-d iatom Coccolithophores phosphatic facies facies (mean 2.2%) mud, laminated Zooplankton: Benthic: Phosphate: Economic deposits: Calcareous Foraminifera bivalves, >20% locally petroleum source, Radiolaria brachiopods- diatomite Benthic: siliceous facies Cd, Cu, Mo, Se, Fe, Zn Bacterial mats ratios like modem plankton Shublik Fm. Triassic, North Glauconitic rare foraminifera Marine reptiles TOC: Slope, Alaska flne ss or sit, Ichthyosaurus, <1-6 wt.% 6-178 m thick 10-50 %glauconite shastasaurids) (mean 0.7-1.8 %) Phosphatic Economic deposits: (sit, sit-ls. Is) + phosphate Bivalves (Halobia sp.. Phosphate: Petroleum source nodules Monotis sp.) in organic-rich <1-14% (mean 0.1-7.8%) Organic-rich facies Is, marl, ms lamins (shelly) Relative enrichments of Co, Brachiopods & molluscs in Cu, Ni in organic-rich phosphatic facies facies; U in phosphatic facies

a\ 00 TABLE 1.6. (continued)

Phosphoria Fm. Middle-Late Permian, ID, Tosi Chert M. acritarchs? (in Bryozoans, brachiopods. TOC; MT, WY, UT ch,sandy ch phosphorites) pelecypods, echinoids-Tosi < 1 -7 wt.% (mean 2.3% in 90-392 m thick Retort Phosphatic Shale M. phosphorite) carb ms, ph Fish, brachiopods. Economic deposits; Cherty Shale M. pelecypods, gastropods- Phosphate: phosphate. cherty ms, ms Retort M. <1-39 wt.% (mean 9.6%) vanadium. Rex Chert M. petroleum source chert; cherty ms. Is, Sponge spicules. Normative calculations and ph inarticulate & articulate suggest marine biogenic Meade Peake Phosphatic brachiopods, crinoids. enrichment in Cu, Mo, Zn Shale M. biyozoans- Rex Chert M. and possibly Cd and Ni and ph, carb ms, (Is) marine hydrogneous Lower Chert M. fish debris, conodonts. enrichment of Cr, V, and ch, (ph) inarticulate brachiopods, REE's bivalves, cephalopods, and gastropods-Meade Peake ^ PInecrest Sand ss. Is not reported gastropods (esp. Turiiella depletion of "C and (and Pliocene units) sp.), bivalves, cetaceans. enrichment of "O in western Florida pinnipeds, seabirds (mass mollusc shells suggests kill assemblage) seasonal high productivity Economic deposits; shell mines many cool-water forms

' compiled from (Allmon et a!., 1996; Barnes, 1977; Barron, 1986; Garrison et al., 1987; Kupecz, 1995; MacKinnon, 1989a; MacKinnon, 1989b; MacKinnon, 1989c; McKelvey et al., 1959; Medrano and Piper, 1994; Parrish, 1986; Parrish, 1987; Piper and Isaacs, 1995; Piper and Medrano, 1994; Pisciotto and Garrison, 1981; Savrda and Bottjer, 1987; Sheldon, 1989) " abbreviations; ss=sandstone; slt=siltstone; ms=clastic mudstone; ls=limestone; ch=chert; ph=phosphorite; carb=carbonaceous; ( )-minor component diatom biostratigraphy (Barron, 1986) present in upper 2/3 of section (MacKinnon, 1989b) ^ inarticulate brachiopods and some fish debris dominate fauna in ph beds; molluscs common in shaly limestones/calcareous shales

ONvO TABLE 1.7. Suggested criteria for classifying sediments as most likely, likely, maybe, unlikely, or most unlikely to represent upwelling environments. See text for discussion and references. TYPE! MOST LIKELY YES LIKELY YES MAYBE LIKELY NOT MOST LIKELY NOT hp.ge ••Si-P-C facies geometry (100-km scale) ORt •Any 2 upwelling-related lithologies (Si+P, *1 upwelling-related Terrestrial organic hp, ±od Si+C, P+C) lithology (Si, P, or Ct) matter Terrestrial organic matter AND/OR AND/OR *Oxygcn-minimum zone edge effects hp, od (sedimentary and biologic) ± OR OR hp High levels of trace elements ± Organic-rich rocks displaced to one side of Organic-rich rocks in the basin basin center ± any of the following: hp, ±od 1. *Organic-rich fecal pellets hp 2. Fish debris§ 3. Abundant planktic and pelagic organisms, including microplankton as hp well as large predators cw 4. Cool-water taxa 5. *Lack of shallow-water reefs but Extensive, shallow-water abundant suspension feeders lacking reefs (based on analogy hp, cw symbionts with Recent reefs)

hp 6. *Seabirds, guano (Cretaceous & younger) • OR OR OR Any 4 or more numbered biological indicators Any 3 biological Any 2 biological listed above indicators indicators {Codes: hp High productivity **known only from coastal, arid areas that experience year-round upwelling ge Geography of upwelling •characteristic of high productivity areas od Oxygen depletion tmarine-source, >0.5 wt % TOC and/or oil-prone cw Cool-water conditions (jcephalopod debris might also be important 71

CHAPTER 2

PALEOCEANOGRAPHY OF THE UPPER TRIASSIC LUNING FORMATION AT

THE SHOSHONE MOUNTAINS, NEVADA 72

INTRODUCTION

As discussed in Chapter I, macrofaunal characteristics of ancient upwelling zones

are not as well-studied as microfaunal and geochemical characteristics. This is somewhat

surprising, given the profound impact of macro-organisms on the sedimentology and

geochemistry of upwelling areas. General characteristics of faunas in nutrient-rich

ecosystems have been summarized by Brasier (1995) and Parrish (1995) (Table 2.1).

Nutrient-rich (eutrophic) ecosystems differ in several important ways from oligotrophic

(normal-marine) ecosystems. Eutrophic ecosystems tend towards abundant, unstable

populations of a few dominant taxa, whereas oligotrophic ecosystems tend towards more

stable populations of many taxa. Mass-mortality commonly occurs in eutrophic systems,

and benthic faunas tend to be depauperate in connection with oxygen-depleted bottom

conditions (Brasier 1995). In eutrophic systems, nutrient abundance can promote

zooplankton blooms to such an extent that organic-rich fecal pellets are deposited and

preserved as sediment rather than scavenged and recycled in the water column (Honjo

and Roman 1978). Eutrophic systems commonly contain non-symbiotic suspension feeders (e.g., bryozoans), whereas oligotrophic systems commonly contain suspension feeders possessing algal symbionts (e.g., zooxanthellate corals) (Brasier 1995). Among eutrophic systems, upwelling zones tend to have taxa indicative of cool waters, even at tropical latitudes (Hallock and Schlager 1986). Each of these characteristics of eutrophic ecosystems has the potential to be preserved in the fossil record. 73

Macrofauna are potentially useful in identifying ancient upwelling areas, especially in cases where more familiar geochemical facies, such as phosphorite, chert, and organic carbon were not deposited or are not preserved. Upwelling and enhanced productivity that results in a primarily biological signal occurs today in the Northwest

African upwelling zone (see Chapter 1). Ecological characteristics of macrofauna in upwelling areas are also potentially useful in evaluating faunas without modem analogs

(see Tables 4 and 5 in Brasier 1995).

Understanding the composition and ecology of macrofauna in upwelling zones also sheds light on variability among upwelling areas and particularly on identifying the range of upwelling intensity in the geologic record (see Chapter 1). Rock units with abundant phosphate, organic carbon, and possibly chert or glauconite are fairly easy to identify as upwelling deposits, for example, the Miocene Monterey Formation and the

Permian Phosphoria Formation. These two formations contain abundant macrofauna, including marine vertebrates (fish; also seabirds and whales in the Monterey Formation,

Yochelson 1963; Barnes 1977; Warheit 1992). These two formations were deposited on a west-facing coast and experienced coastal upwelling, similar to modem upwelling zones.

The situation becomes more complicated when dealing with depositional conditions without modem analogs, for example upwelling in epicontinental seaways

(Parrish 1982). The Sharon Springs Member of the Pierre Shale contains abundant phosphate and organic carbon and a rich biota of marine vertebrates (Parrish and Gautier 74

1993). This unit has been interpreted in contrasting ways, as resulting from either

deposition in a salinity-stratified basin or from enhanced productivity and upwelling.

Parrish and Gautier (1993) argued that displacement of facies to one side of the basin,

geochemical and faunal evidence for depauperate benthos, and abundant vertebrates,

phosphate, and organic-rich fecal pellets support an upwelling origin for the Sharon

Springs. In this case, sedimentological, geochemical, and paleontological evidence—in

short, all available data—were used to support this conclusion.

In other cases, a rock unit was deposited in a paleogeographic position amenable

to upwelling but contains few geochemical indicators of upwelling. The Pinecrest Sand

and other Pliocene units in southwestern Florida contains abundant and diverse

molluscan and vertebrate assemblages with both cool- and warm-water affinities (Allmon

1993; Allmon et al. 1996). The cool-water forms are present in great abundance during

specific time intervals interpreted as periods of high productivity (Allmon et al. 1996).

The sediments consist largely of shells and a little clastic material. Phosphate is abundant

in beds below the Pliocene units, but phosphate, organic carbon, or siliceous sediments

are lacking in the Pliocene units (Allmon 1993; Allmon et al. 1996). Similarly, there is

no evidence for depauperate benthos in these nearshore sediments, and offshore equivalent facies that might hold evidence for the oxygen-minimum zone are not exposed. Coastal or topographic upwelling could have occurred along the southwest coast of Florida in the Pliocene (Alknon et al. 1996). Several lines of paleontologic evidence, including faunal abundance, cool-water C and O isotope signatures firom 75

mollusc shells, presence of isolated coral colonies indicative of cool, turbid

environments, lack of calcareous algae, and presence of cool-water forms that no longer

inhabit the area suggest that upwelling enhanced productivity and consequently affected

the sedimentary record (Allmon et al. 1996).

The examples explained above illustrate the potential utility of macrofauna in

identifying and characterizing ancient upwelling zones. In this chapter I discuss results

of a paleoceanographic study of the Luning Formation, a rock unit that contains an

abundant biota and that was deposited along the west coast of North America during the

Late Triassic.

The Luning Formation in Nevada is famous for large (ca. 15 meters long)

ichthyosaur fossils of the genus Shonisaurus. Remains of 37 or more ichthyosaurs are

found at Berlin-lchthyosaur State Park (BISP) alone (Camp 1980). Camp (1980) and

Hogler (1992b) each observed that ichthyosaur fossils in the Shoshone Mountains at

BISP occurred throughout the marine part of the section, implying that the animals lived

in the area for an extended period of time. Additionally, Hogler (1992b) found evidence for a mass-mortality event involving at least 9 ichthyosaurs, indicating local abundance.

Hogler (1992b) suggested that upwelling could generate a rich food source that would attract ichthyosaurs. Parrish and Parrish (1983) suggested that Mesozoic marine reptiles could have filled an ecological niche in upwelling zones similar to whales today.

Large ichthyosaurs would likely need abundant food, and it is therefore reasonable to 76

predict tiiat they would gather in prey-rich upwelling zones. The Luning Formation contains abundant macroinvertebrates that could have served as prey.

This study examines the hypothesis that the Luning Formation at the Shoshone

Mountains was deposited in an upwelling zone. The Luning Formation was deposited in a back-arc basin which could have experienced upwelling conditions (Parrish and Curtis

1982). To conduct this study, I examined the petrology, geochemistry, and paleontology of the Luning Formation in the Shoshone Mountains, Nevada. At this locality, outcrops of the Luning are the best-studied in terms of both sediments and fossils (Silberling

1959; Hogler 1989; 1990; 1992a). The stratigraphic succession at BISP is fairly complete and is the least structurally complicated of the Luning localities. 77

TECTONOSTRATIGRAPHIC SETTING

Mesozoic rocks of the western Great Basin consist of packages of fault-bounded marine carbonate, clastic, and/or volcanogenic rocks. The fault-bounded packages are collectively termed the Mesozoic marine province (MMP; usage following, for example.

Speed 1978b; Oldow 1984). The Late Triassic Luning Formation is part of one of these packages, and it outcrops in the southeastern part of the MMP. Current interpretations favor the deposition of MMP rocks at low latitudes (within 15° of the paleo-equator,

Tozer 1982) in a back-arc basin on the western edge of Pangea (Figure 2.1). A subduction-related, magmatic arc lay to the west and the craton lay to the east (Speed et al. 1989; Oldow et al. 1993; Lawton 1994). Deposition in the MMP began after the Early

Triassic Sonoma Orogeny and ended in the late Early Jurassic, as represented by the synorogenic Dunlap Formation (Lawton 1994). Several tectonic episodes have fragmented, deformed, and metamorphosed MMP rocks, including the Early Cretaceous

Sevier Orogeny (Lawton 1994), late Mesozoic strike-slip displacement (Oldow et al.

1993), and late Cenozoic Basin and Range deformation (Lawton 1994).

Deformation from these events hampers lithofacies correlations across the basin and also from the basin to the craton and adjacent rocks. Today, the Luning-Fencemaker thrust belt underlies rocks of the MMP and leaves the Paleozoic and earliest Mesozoic geology largely unknown. Poorly exposed basement rocks, widespread deformation, and disparate but adjacent lithologies have resulted in differing views on MMP formation, especially of basinal rocks, which have thrust faults as basal contacts (e.g.. Speed 1978b; 78

Oldow 1984; Silberling et al. 1987; Speed et al. 1989; Silberling 1990; Oldow et al. 1993;

Lawton 1994). Silberling et al. (1987) and Speed et al. (1989) cautioned that differences

among MMP lithologic successions suggest different depositional and displacement

histories, with each succession an accreted terrane and subsidence occurring via back-arc

spreading. Speed et al. (1989), however, argued that MMP rocks share a common

basement (Sonomia, remains of a Paleozoic arc, Figure 2.2.) that collided with North

America in the Early Triassic, and that subsidence occurred via cooling of Sonomia's

magmatic crust.

Large-scale lithologic patterns suggest that MMP rocks were deposited together and later dismembered and displaced, regardless of the nature of the basement. MMP

lithologies can be grouped into large-scale assemblages from east to west (Speed 1978b;

Speed et al. 1989; also see reconstructions in Lawton 1994): shelf carbonates and clastics, slope carbonates and turbidites, basinal carbonates and clastics, basinal volcaniclastics and carbonates, and arc-related volcanics. This succession of large-scale lithofacies, though unrestored, suggests that the MMP deepened to the west and also supports its origin as a coherent paleogeographic and depositional assemblage. Major thrust and strike-slip faults later juxtaposed assemblages (for example, basinal rocks are thrust eastward onto eastern shelfal carbonates).

The arc on the western border of the MMP erupted intermittently during the

MMP's existence, and consisted of island chains to the north and a continuous chain of volcanoes to the south (present-day southwest U.S. and Mexico, Figure 2.1) (Silberling 79

and Wallace 1969; Lucas and Marzolf 1993; Marzolf 1993; Riggs and Blakey 1993;

Saleeby and Busby 1993; Schweickert and Lahren 1993; Wyld and Wright 1993). Rocks

in the western part of the MMP are predominantly volcaniclastic. Upper Triassic rocks in

the eastern MMP do not contain identifiable volcanic material, suggesting that either the

arc was quiescent during that time or that any volcanic material in eastern MMP rocks

has been altered beyond recognition.

On the eastern side of the MMP, provenance and paleocurrent studies have

established links to the craton for the Late Triassic. To the east, shelfal rocks of the

MMP are separated from continental Late Triassic rocks by several hundred kilometers of

missing record or non-deposition (Lawton 1994). Within this area, one outcrop near

Currie, Nevada, was assigned to the terrestrial Chinle Formation (Lupe and Silberling

1985). The Chinle Formation is famous for spectacular petrified wood. Blocks of

Auricaryoxylon wood are found in clastic sediments of the Dun Glen Formation in the

Stillwater and Tobin Ranges (Silberling and Wallace 1969), suggesting a tie with cratonic

fluvial-deltaic systems (Silberling 1959). Silberling and Wallace (1969) suggested that

terrigenous clastics of the Late Triassic Auld Lang Syne Group (northwest part of the

MMP) were transported firom paleodrainages on the craton. Subsequently,

paleodrainages in the cratonic Chinle Formation were shown to have drained

northwesterly, into the MMP (Lawton 1994, and references therein). Lupe and Silberling

(1985) linked fluvial-lacustrine cycles in the Chinle with episodes of clastic deposition in

the Auld Lang Syne Group. Sandstones firom both the Currie, Nevada, Chinle locality 80

and the clastic Osobb Formation of the Auld Lang Syne Group contain a distinctive

population of detrital zircon grains that were likely derived from plutons in the Wichita

Mountains. This finding provides further evidence for a link between the craton and at

least the northern part of the MMP during the Late Triassic (Gehrels and Dickinson

1995).

Farther south, several lines of evidence suggest a less far-traveled depositional tie

between the Luning Formation and the North American craton. Chert-pebble

conglomerates, sands, and argillites represent deltaic settings close to a high-relief source

(perhaps fan-deltas) in the Pilot Mountains (Oldow 1981). Chert-pebble provenance

appears to be from the upper Paleozoic Mina Formation (part of either Sonomia or the

accreted Golconda allochthon. Speed et al. 1989) and not from more inboard continental

sources. Rare, small wood fragments have been found in argillites of the lower, clastic

member of the Luning in the Shoshone Mountains (Silberling 1959). Reilly and others

(1980) described two petrographic provinces from sandstones at three Luning Formation

localities of different ages. The provinces included a quartz, sedimentary-lithic province

typical of older outcrops to the northeast, and a feldspar, volcanic-lithic province typical

of younger outcrops to the southwest. They suggested that the west-east pattern of decreasing volcanic material roughly mirrored the original west-east geographic relations among these localities(Reilly et al. 1980). More recently, single-grain detrital zircon analysis of sandstones from the early Norian part of the Luning in the Pilot Mountains 81

(Nappe 2d of Oldow 1981) yielded zircons with both arc and cratonic affinities

(Manuszak et al. 1997).

In this chapter, I will assume that the Luning Formation was deposited on the eastern side of a back-arc basin on the western margin of North America. Regional lithologic distribution, zircon analyses, provenance studies, and the distribution of fossil wood all support this interpretation. 82

LITHOLOGY, AGE, AND PALEONTOLOGY OF THE LUNING FORMATION

The Luning Formation is a mixed carbonate-siliciclastic unit that varies in

lithology and age among localities (Figure 2.3). The formation contains impure

carbonates and shales, clastic rocks, and limestones and dolomites and varies in thickness

from 1-2.8 km. First, I describe the facies found at the Shoshone Mountains section, and

then I briefly describe sections at other localities.

At the Shoshone Mountains. Silberling (1959) divided the Luning Formation into

four informal members: clastic, shaly limestone, calcareous shale, and carbonate (Figure

2.4). Contacts between the members are gradational. The clastic member is about 200

meters thick and contains very thick-bedded conglomerate in its lower half. The base of

this member lies in broad channels eroded into limestone of the Middle Triassic

Grantsville Formation. The middle part of the clastic member contains interbedded and

interfmgering conglomerate, poorly sorted, greenish-gray, cherty sandstone with interstitial sericite, and lenses of brown to green argillite. The upper 35 meters of the member contain mostly highly fractured argillite with a few thin sandstone interbeds.

Beds are largely structureless. Rare petrified wood {Araucarioxylon?, in conglomerate) and a twig impression {Palissya, in argillite) are the only fossils known from this member. The top 8 meters of the clastic member grade into the shaly limestone member

(Silberling 1959).

The shaly limestone member is about 220 meters thick and contains muddy, silty, brownish-gray wackestones and packstones with interbedded noncalcareous to calcareous 83

shale. The proportion of shale relative to limestone increases upsection. No primary sedimentary structures are visible and bioclasts do not show preferred orientations.

These rocks contain abundant benthic pelecypods, especially in the lower 1/3 of the member. Other fauna include gastropods, ammonoids, echinoderms, nautiloids, rare crustaceans, and ichthyosaurs (Silberling 1959; Hogler 1989; 1992a).

The proportion of carbonate increases upsection in the shaly limestone member, with argillite gradually giving way to calcareous shales of the calcareous shale member.

Silberling (1959) recognized this member by its purplish-red weathered color. This member is about 190 meters thick and contains fine-grained, purplish-weathering shales and thin limestones. Limestone beds increase in both thickness and proportion upsection.

Fauna include thin-shelled pelecypods and cephalopods. Other benthic fauna are absent

(Silberling 1959; Hogler 1989; 1992a).

Mud and silt content decrease into the carbonate member. Beds of this member are the best exposed of all Luning marine rocks and form a prominent bluff on the southeast wall of West Union Canyon. The upper half of this unit is covered and/or faulted out near BISP, but contains limestone and algal-laminated dolomite (Whitebread et al. 1988) where it is exposed farther south in the Shoshone Mountains. In West Union

Canyon, the carbonate member (lower half) is about 325 meters thick and consists of fairly pure carbonate, some secondary dolomite, and minor shale interbeds (middle 1/3 of the member). Fauna include brachiopods, echinoderms, pelecypods, gastropods, and rare ichthyosaur fragments (Silberling 1959; Hogler 1992a). 84

In the Pilot Mountains (Figure 2.3), the Luning Formation consists of the

following members from oldest to youngest: a lower shaly limestone member, a middle

clastic member, and an upper carbonate member. In its type locality in the Pilot

Mountains (nappe N4, Muller and Ferguson 1936; Oldow 1981), the Luning is

continuous and 2.5 km thick (lower member 900 m, middle member 900 m, upper

member 750 m). The shaly limestone fauna differs from that at BISP; in the Pilot

Mountains, Stanley (1979) found 5-6 successions of small-scale coral and sponge

buildups in the lower member. In the Pilot Mountains, the middle member contains

conglomerates and sands with up to 60% feldspar sand in a few outcrops, suggesting

proximity of a volcanic source (Oldow 1981). Carbonate rocks in the upper member are

fairly pure and contain dolomite (Oldow 1981), as at BISP. Luning rocks in the Pilot

Mountains are not associated with older rocks but instead are underlain by thrust faults

(Oldow 1981). The Luning section at the Gabbs Valley Range (Figure 2.3) consists of

limestones, subordinate shales, and an upper dolomite sequence (Muller and Ferguson

1936; Oldow et al. 1993) that is roughly equivalent to the upper member of the Luning in

the Pilot Mountains.

Stratigraphic sections of the Luning Formation at the Paradise Range are more similar to the BISP section, whereas sections in the Cedar Mountains are more similar to

the Pilot Mountains succession (Speed et al. 1989). At the Paradise Range, west of BISP, clastic rocks grade into basinal and then shelfal limestones and finally dolomite, but volcanic components dominate the lower part of the section and include volcanic breccia 85

and mafic intrusive and extrusive rocks. Thicknesses have not been measured because of severe deformation and metamorphism. In the southern Cedar Mountains, southwest of

BISP, there are no volcaniclastic rocks, but a succession of red beds and evaporites, probably in fault contact, is exposed between calcareous shales and purer limestones and dolomites. These differences highlight the difficulty of correlating among localities

(Speed et al. 1989). The base of the Luning is in depositional contact with underlying rocks only at the Shoshone Mountains, where the Middle Triassic Grantsville Formation unconformably underlies the Luning (Silberling 1959). At all other localities, however, the Luning lies in fault contact with underlying units (Oldow 1981).

The clastic facies of the Luning appears to represent marginal to shallow-marine deposition. In Luning outcrops in the Pilot Mountains, variability in thicknesses and positions of chert-pebble conglomerate, sandstone, and shale resulted from the introduction and migration of a fan-delta into the marine basin (Oldow 1981). A variety of sedimentological and paleontological evidence favors interpretation of the clastic member as deltaic rather than deep-marine in origin. In the Shoshone Mountains, carbonates of the Middle Triassic Grantsville Formation are separated fi*om the overlying clastic member of the Luning Formation by an irregular, karsted surface. This surface was probably formed at least partially by channelization, as a few broad, shallow channels overlie the surface and contain Grantsville carbonate clasts in addition to

Luning clastic sediments (Silberling 1959). The disconformity implies subaerial exposure between Luning and Grantsville deposition, and a deltaic origin for the Luning 86

clastics would require less of a base-level drop than a deep-marine origin. Marine

microfossils have not been reported from Luning clastics, but twig impressions and wood

fragments are present occasionally in the Shoshone Mountains section (Silberling 1959).

Thick packages of clastics in the middle member of the Luning in the Pilot Mountains

were likewise interpreted as representing delta-plain to pro-delta environments by Oldow

(1981). He explained a lack of marginal marine fossils in clastics as resulting from a

high-gradient delta system draining a high-relief source area (Oldow 1981).

At all localities, the Luning Formation shallows upward to carbonate-dominated

lithologies, shown by the succession from clastics or shaly limestones to purer limestones

topped by dolomitic, algal-laminated beds that occur at the top of all Luning sections

(Speed etal. 1989; Figure 4A in Oldow et al. 1993). Overall, the Luning Formation

reflects rising sea level (Silberling 1959; Oldow et al. 1993). Oldow and others (1993)

interpreted the upper member of the Luning Formation as representing the culmination of

shallow-marine carbonate deposition, with the uppermost dolomite representing

progradation into intertidal marine environments. The dolomite represents their sequence

boundary 2 and is between late early Norian and late late Norian in age (Oldow et al.

1993). Whereas the top of the Luning Formation is everywhere similar in age, lower

members of the Luning Formation vary in age across localities, from late Camian to early

Norian (Silberling and Roberts 1962; Silberling and Tozer 1968; Kristan-Tollmarm and

Tollmann 1983) (Figure 2.3). 87

Silherling (1959) reconized 3 ammonoid zones, each about 30 meters thick, in the

Luning Formation at BISP. These zones are the Klamathites schucherti zone (middle

upper Camian), the Klamathites macrolobatus zone (upper upper Camian), and the

Guembelites zone (lower Norian) (Silherling 1959; Silherling and Tozer 1968).

Occurrences of spiriferid and terebratulid brachiopods in strata above these zones

indicate an early to early middle Norian age {Mojsisovicsites kerri and Juvavites magnus

zones, Silherling and Tozer, 1968; Sandy and Stanley, 1993). Other fossils at BISP hold

promise for biostratigraphic refinement, notably conodonts and halobiid bivalves, both of

which occur in the calcareous shale member.

In the Pilot Mountains, low-relief coral-sponge buildups near the top of the lower

member were assigned to the early Norian based on the presence of Mojsisovicsites kerri

(Silherling, cited in Sandy and Stanley 1993). The occurrence of Paracestes in the same

member also indicates an early Norian age (Kristan-Tollmann and Tollmann 1983). The

base of the upper member contains Indojuvavites and ammonoids characteristic of the lower middle Norian Juvavites magnus zone (Oldow 1981). The Luning in the Pilot

Moimtains spans at least the early-middle Norian. Luning deposition in the Pilot

Mountains thus began slightly later than deposition in the Shoshone Mountains.

Invertebrate macrofossils known from the Luning at all localities include pelecypods, cephalopods, gastropods, sponges, corals, hydrozoans, and articulate brachiopods; microfossils include calcareous foraminifera and conodonts (Silherling

1959; Stanley 1979; Kristan-Tollmarm and Tollmann 1983; Hogler 1990; 1992a; 1992b; 88

Silberling, personal communication, 1996; this study). Most of the shaly limestones in the Luning Formation contain benthic mollusc fossils, while calcareous shales often contain ammonoids and thin-shelled bivalves (Muller and Ferguson 1936). Among the vertebrates, ichthyosaurs and rare fish {Ceratodus, Hogler, 1992a) are known. Almost all ichthyosaur fossils are members of the genus Shonisaurus, one of the largest known ichthyosaurs (Kosch 1990; McGowan 1996a). Ichthyosaur fossils are found in the lower member of the Pilot Mountains (Stanley 1979), basinal limestones of the Cedar

Mountains (D. Montague-Judd, unpublished field notes), and the shaly limestone at BISP

(Camp 1980; Hogler 1992b). Only the ichthyosaur fossils at BISP are well documented.

In the 1950s and 60s, Dr. Charles Camp excavated the partial remains of about 37 ichthyosaurs of the genus Shonisaurus from a single canyon in the Shoshone Mountains

(Camp 1980), making BISP North America's richest ichthyosaur deposit (Kosch 1990). 89

METHODS

The goal of this study is to analyze the Luning Formation in terms of upwelling indicators (see Introduction and Chapter 1 for discussion of indicators). I used sedimentological, geochemical, and paleontological data to conduct this analysis.

This study focused on Luning Formation outcrops in the Shoshone Mountains in and around BISP. I measured, described, and sampled 5 stratigraphic sections from all of the marine Luning strata exposed in West Union Canyon (Figure 2.4). The marine strata include all of Silberling's (1959) shaly limestone and calcareous shale members, and the lower 1/2 of his carbonate member. The upper part of Silberling's (1959) carbonate member is not exposed in this canyon. It is exposed farther south, but is severely metamorphosed and folded, with neither the top nor the bottom of the member exposed

(Silberling 1959).

Ninety slabs, 33 standard thin sections, and 39 acetate peels were made for petrographic analysis. Acetate peels were made by polishing slabs with 600 grade silicon carbide grit, etching in 10% HCl, and transferring the etched surface with acetone onto 5 mil acetate film.

Rock specimens for geochemical analysis were chosen to represent each group of lithologies represented in the section (e.g., calcareous shale, silty limestone). I chose the least weathered hand samples for the analyses and crushed samples into chips that were then sent to the laboratory for grinding and analysis. About 2 grams of each sample were sent for analysis. DGSI Total Quality Geochemistry (The Woodlands, TX) analyzed 36 90

samples for total organic carbon (TOC) content by combustion in a LECO carbon analyzer. Samples approaching 1 weight percent TOC were analyzed by Rock-Eval pyrolysis. Rock-Eval determines the amount of free hydrocarbons and their maturity in the rock sample. XRAL Laboratories (Toronto, Ontario) analyzed 37 samples for major and minor elements, using agate milling, HF/HNO3/HCIO4 extraction, and induction- coupled plasma optical emission spectrometry (ICP-OES).

Trends in faunal and taphonomic data can reveal high-productivity and/or low- oxygen environments that are often associated with upwelling (Mullins et al. 1985;

Allmon et al. 1995). I used field observations of macrofossils, petrographic observations of microfossils and biofabric, and Hogler's (1992a) faunal lists to evaluate whether or not oxgyen-minimum-zone edge effects were present in the section. I also used these data to assess whether or not faunas resembled those in other rock units that have been interpreted as upwelling environments, such as the Late Triassic Shublik Formation

(Parrish 1987a) and the Permian Phosphoria Formation (Wardlaw and Collinson 1986).

Key criteria for recognizing high-productivity faunas include presence of cold-water forms (and absence of warm-water forms and features) and low-diversity, high- abundance assemblages (see Chapter 1 for further discussion of biological indicators of upwelling zones). 91

RESULTS

Lithofacies Descriptions and Interpretations

Appendix A contains the 5 measured stratigraphic sections, which together represent a composite Luning section of about 700 meters (Figure 2.5). Covered intervals and faulting precluded exact bed-by-bed correlation, but several laterally continuous marker beds are identifiable in the section (Figure 2.6). Most of the samples for thin sections and geochemical analysis were from limestones, since most of the shales were highly weathered and poorly exposed.

I estimated sedimentation rates in the Luning at BISP for biostratigraphic zones a, b, and c of Silberling (1959) (Table 2.2). Ignoring compaction and dissolution, these calculations yielded rates of 1.5-2.7 cm/1000 year for three ammonoid zones in portions of the Sla, Sbl-Scs, and Scs-Pel subfacies (see descriptions below).

Petrographic study revealed trends in sorting, texture, fossil preservation, and diagenesis that follow major lithologic changes in the marine part of the section. These trends were consistent with previous work (Silberling 1959: Hogler 1989; 1992a).

Below, I describe new lithofacies observed in the current study. These lithofacies emphasize sedimentary textures and are more detailed than those of Silberling (1959) or

Hogler (1992a).

Lithofacies were based on field and petrographic observations of lithology, texture, sedimentary structures, bedding, taphonomic features, and fossil content. The 92

lithofacies include shaly, fossiliferous limestone (S), pel-bioclastic limestone (P), oolitic bioclastic limestone (O), and fossiliferous limestone (F) (Table 2.3, Figures 2.7- 2.13).

Shalv. Fossiliferous Limestone Facies (S)

Description

Lithologies of this facies occur in the lower half and uppermost 50 meters of the composite section and include packstones, wackestones, and shales with varying amounts of fine-grained clastic material, whole fossils, and bioclasts. Three subfacies comprise this facies: silty, brown limestone and argillite (Sla), silty, black limestone (Sbl), and calcareous shale (Scs). Ichthyosaur bones are common in thin wackestone ledges of the

Sbl subfacies.

Siltv. brown limestone and argillite subfacies fSla. Figure 2.7). All of the limestone beds have hackly-weathering, tan, brown, gray, and green surfaces with no primary sedimentary structures visible. Beds range from 0.5-5.5 meters thick, with most limestone beds about 1 meter thick. Wackestones are thickest and whole-fossil packstones are thinnest. Silty, bioclastic packstones occur at the base of the marine part of the section and overlie sands and argillites of the clastic member. The packstones grade into whole fossil wackestones, which are then succeeded by very silty whole fossil packstones and finally bioclastic wackestones with scattered whole fossils. The bioclastic packstones and whole fossil wackestone beds gradually thicken and thin laterally, while the silty, whole-fossil packstones are discontinuous. 93

In terms of texture and fossil content, the bioclastic packstones at the base are poorly sorted and contain many echinoderm and bivalve fragments. Bioclasts do not show a preferred orientation and are densely packed. The proportion of whole fossils

(bivalve molluscs in particular) increases upsection to the very silty packstones, which are very poorly sorted and contain abundant benthic pelecypods and gastropods and rare ammonoids and nautiloids. Thin (< 1 meter) patches of densely packed Palaeocardiata sp. or Lopha cf. montiscaprilis (Klipstein) (C. McRoberts, personal communication,

1998) occur in the very silty packstones. Fossils occur in both dense patches and loose aggregations and tend to have their long axes oblique or parallel to bedding in convex- down and convex-up orientations. Whole fossils, disarticulated shells, and hash are present, and fragments range from angular to rounded. Some of the limestones in this facies show mottling on weathered faces, but burrows are not visible on slabs or in thin section.

Both the fossils and the micritic carbonate matrix of the limestones are recrystallized, and pressure solution seams are common. Fossil preservation types include internal molds (bivalves, gastropods), recrystallized hard parts with mud and sparry calcite fills (whole bivalves, cephalopods), syntaxial overgrowths (echinoderms, bivalves), and rare impressions. Most gastropods observed contain mud fills of darker color than the surrounding, recrystallized matrix.

Thick, brown, brown-gray, or greenish-brown argillites are interbedded with the limestones and comprise about 40% of the subfacies thickness. Individual beds range 94

from 1 to 5 meters thick. The argillites are weathered, buried, non-calcareous, and non-

fossiliferous except for rare whole fossils (only where interbedded with whole fossil

packstones) and occasional shell-packed, calcareous nodules. The nodules contain

monotypic clusters of recrystallized, disarticulated, thin-shelled bivalves, including

1Halobia sp. (at the base of the marine part of the section) or compacted, whole and disarticulated, ostreid bivalves (in the rest of the subfacies). Smaller shells and

fragments, such as Halobia sp., are aligned parallel to subparallel to bedding, while larger shells, such as ostreid bivalves, are aligned more obliquely and more randomly to

bedding.

Siltv. black limestone subfacies (Sbl. Figure 2.8). The silty, black limestone subfacies includes two main types of beds: 1) thick-bedded silty micrites and wackestones, and 2) thin ledges of fossiliferous packstone and wackestone. Limestones of the Sbl subfacies are interbedded with calcareous shales (sub-facies Scs below).

The thick-bedded silty micrites and wackestones weather tan or blue-gray and have black fresh surfaces. Beds are prominent, continuous, and 2-5 meters thick.

Primary sedimentary structures are not visible and numerous fractures and calcite veins are present. A few beds have randomly oriented structures visible on weathered surfaces that resemble burrows. These structures are centimeter-scale blobs or patches that weather orange or gray, have sharp contacts with the surrounding matrix, and contain small bioclasts. Fossils commonly found in these beds include scattered whole gastropods, pelecypods (disarticulated, including thin-shelled forms), and ammonoids. 95

Micrites contain iiighly dispersed, randomly oriented, whole fossils and single shells.

Wackestones contain dispersed fossils and scattered patches of loosely packed fossils, particularly small gastropods. In the lowermost occurrences of the wackestone beds, from the 90-100 meter level in the composite section, the proportion of fragmented to whole fossils increases within each bed. Bivalve shells show convex-down and convex- up orientations. Bioclasts tend to be poorly sorted and oriented concordantly or obliquely to bedding.

The thin ledges of fossiliferous packstone and wackestone weather blue-gray to tan and are dark gray on fresh surfaces. The beds are about 0.5 meter thick and occur over a 30-meter stratigraphic interval beginning at about the 100-meter level in the composite section. Primary sedimentary structures are absent, but cm-scale burrow mottling is visible in slabs and thin sections. Ledges contain patches of small, thin, recrystallized bivalve shells (articulated and disarticulated), small gastropods, ammonoids

(including juvenile sizes), and rare echinoderm fragments, ostracods, and foraminifera.

Ichthyosaur bones are common as float and in the ledges and include partially articulated skeletons and isolated bones. Partially articulated skeletons occur in discontinuous patches scattered laterally and vertically through exposures of these ledges. Bones are abraded even when in partial association, but small, rounded bone pebbles are absent.

Invertebrate fossils are heterogeneously distributed in the rock, and fossil patches are densely to loosely packed in a mud matrix. Fossils are very poorly sorted and orientation is variable, although many single shells have their long axes oblique to bedding. Shell 96

fragments are common, disarticulated shells less conrunon, and articulated, bivalved animals rare (bivalve molluscs and ostracods). A few beds contain moderately sorted, disarticulated nuculid bivalve shells of medium thickness oriented obliquely to bedding.

Ammonoids are oriented with the axis of coiling normal to bedding (whorls are parallel to bedding).

Among both main bed types, the matrix is brown, recrystallized, and has a clotted appearance in thin section. Calcite veins, pressure solution seams, and clusters and cubes of oxidized iron (replacing pyrite?) are common. Bioclasts are recrystallized.

Ammonoids are preserved as recrystallized shells filled with spar in the inner chambers and mud in the outer chamber. Gastropods are preserved as internal molds with a darker- colored fill than that of the surrounding matrix, particularly in the lower beds within this subfacies. In the uppermost occurrences of this subfacies, ammonoids (including juvenile sizes), thin-shelled bivalves (Halobia sp.), and/or ostracods dominate the fossil content of the rocks. A few beds contain laminations composed solely of compacted, thin-shelled bivalves.

Calcareous shale subfacies (Scs. Figure 2.9). This subfacies contains poorly exposed beds of tan to purplish-weathering, black calcareous shale with no visible sedimentary structures. Millimeter-scale rounded grains (probably weathered bioclasts) weather out on the surface of the rock. The Scs subfacies is interbedded with the Sbl subfacies (see description above), with beds 0.5 - 5 meters thick. 97

Fossil abundance varies in Scs subfacies beds. Most beds contain dispersed or

loosely packed thin-shelled bivalves and/or ammonoids. A few beds contain densely

packed halobiid bivalves {Halohia cf beyrichi [Mojsisovics]) oriented parallel to

bedding. Other beds appear barren in the field, but contain abundant ostracods, very

small, thin-shelled bivalves, and occasional larval ammonoids in peel and thin section.

Rare sponge spicules and ?radiolaria are also visible in thin section. Halobia cf. beyrichi

(Mojsisovics) valves are disarticulated and preserved in equal numbers in convex-up and

-down orientations (Table 2.4). Only one of the beds examined petrographically actually

contained no macrofossils or microfossils.

The matrix and bioclasts are strongly aligned parallel to bedding, and fossils are

medium-sorted and recrystallized or preserved as impressions. The matrix is dark brown,

recrystallized, and has a clotted appearance. Oxidized iron cubes and clusters are

abundant.

Interpretation

Limestones and shales of the S facies as a whole represent shelf environments

below fair-weather wave base. The presence of fine-grained material and lack of primary sedimentary structures in all lithologies indicate a lack of current winnowing under normal conditions. Terrigenous silt content is highest in rocks of the S facies.

Rocks of the silty, brown limestone and argillite subfacies contain fossil concentrations that suggest episodic deposition by storm events. The lack of primary sedimentary structures suggests that storm deposition was limited to the distal reaches of 98

offshore flow (distal tempestites, Aigner 1985) and resulted in net deposition rather than

erosion. Articulated, spar and mud-filled bivalves (ostreids and Palaeocardiata sp.)

suggest rapid burial of articulated shells (Kreisa 1981). The presence of articulated,

disarticulated, and fragmented fossils in oblique, convex-up and -down orientations

suggests transport and settling of bioclasts and fine-grained material into the depositional environment without further current winnowing. The fossil assemblage then contains

both allochthonous (fragmented) and parautochthonous (articulated) elements (Brett and

Baird 1993). Gastropods filled with mud that differs from the matrix were probably transported into the depositional environment (Horowitz and Potter 1971). Alternatively the mud within the shell might have escaped recrystallization, thereby appearing darker because it is finer-grained, but this is unlikely since the gastropod shells themselves are recrystallized.

Shelly nodules in the argillites represent colonization during background sedimentation. The change from mostly bioclastic packstones at the base of the section to mostly articulated-fossil packstones in the upper half of the Sla subfacies could indicate either lack of whole fossils in lower packstones (the discontinuous nature of storm deposits, Kreisa 1981) or a slight deepening of the basin. The parautochthonous component of the fauna consists of a variety of benthic infaunal and epifaunal animals, suggesting bottom conditions favorable to a range of organisms (Biofacies 1 of Hogler

1992a). 99

Overall, the lower half of the silty, black limestone subfacies records quiet deposition of fine-grained, less-terrigenous material. Fossils are dispersed and occur without a preferred orientation. The 90-100 meter interval in the composite section when fragmented fossils increased in abundance could have been to related to increasing energy conditions, a decrease in net sedimentation rate, or both. Either of these events would have allowed bioclasts to be abraded. The varying orientation of the fossil fragments and the nature of overlying sediments suggest slightly longer exposure on the seafloor (see below), but without much current winnowing. Fossil fragments could have been transported into the area by storms and then exposed on the seafloor for a short time. The dominance of halobiid bivalves and ammonoids and lack of benthic organisms in the upper 100 meters of the Sbl subfacies suggests inhospitable benthic conditions

{Halobia and Mojsisovicsites-Guembelites communities in Biofacies 2 of Hogler 1992a).

Horizons containing abundant ichthyosaur bone reflect longer exposure on the seafioor than other lithologies in the facies. This interpretation is based on the abraded condition of ichthyosaur bones, the abundance of ammonoids, and the presence of fi-agmented and disarticulated shells and disseminated pyrite (Brett and Baird 1986), as well as the presence of possibly transported gastropods. However, evidence for condensation such as burrowed hardgrounds, corroded shells, phosphatic concretions, bone gravel (Norris 1986), and mixing of biostratigraphic zones (Kidwell 1993) is lacking. Bones do not occur uniformly and are not confined to any particular horizon.

Additionally, ichthyosaur bones display similar states of preservation, encrusters are rare. 100

and fragile, thin shells are preserved along with fragmented shells. Many shells are obliquely oriented to bedding but others are in different orientations, and rocks are mud- supported and contain microfossils, arguing against current winnowing as a dominant force shaping these sediments. These beds most likely represent multiple-event concentrations and within-habitat, time-averaged assemblages (Kidwell 1991; Kidwell and Bosence 1991). Benthic fossils are less diverse in these rocks, suggesting less favorable bottom conditions (Tropites and Nuculana communities in Biofacies 2 of

Hogler 1992a). The slightly abraded condition of ichthyosaur bones, as well as the presence of rare encrusters on bones suggests that the substrate was soft but not soupy

(which would have excluded encrusters altogether; Martill 1993).

The calcareous shale subfacies represents quiet deposition in waters below storm wave base. Benthic fossils are least diverse in this subfacies compared to all other facies in the section. Ostracods and halobiid bivalves are the only possible benthic dwellers preserved in sediments of this subfacies. Halobiid bivalves are often the only macrofossils found in rocks of this subfacies. Their disarticulated state argues against in situ burial fi-om an epibenthic habit (differing interpretations in Hogler 1992a; McRoberts

1997). The preservation in almost equal numbers of convex-up and -down orientations would support an epibenthic habit if adjacent, disarticulated shells are matched.

However, of 58 rock surfaces examined, only 4 possible matched shell pairs were observed out of 441 shells counted (Table 2.4). A lack of other current-activity indicators leaves the possibility that the halobiids were pseudoplanktonic, planktonic, or nektonic 101

and settled out of the water column onto the sediment surface (Jefferies and Minton 1965;

Flugel 1982; McRoberts 1997). Kuhry and Kuhry et al. (1975; 1976) suggested that

grainstones of thin-shelled bivalves represent transport in a higher-energy environment,

but the lithologies here contain mud and most shells were probably not transported.

Similar halobiid taphofacies occur in the Late Triassic Shublik Formation of Alaska,

where bedding planes of dense, uncorroded, complete but disarticulated shells suggested

a non-benthic habit (J.T. Parrish, M.L. Droser, and D.J. Bottjer, personal communication,

1998). In both the Shublik and the Luning Formations, bedding planes consisting wholly

of halobiid bivalves (in both the calcareous shale and the silty black limestone subfacies

in the Luning Formation) suggest opportunistic "blooms" of these bivalves in response to

favorable water-column conditions.

Sediments in the Scs subfacies reflect dysoxic and possibly anoxic benthic conditions {Halobia and Mojsisovicsites-Guembelites communities in Biofacies 2 of

Hogler 1992a). Rocks of the Scs subfacies typically contain ostracods even when other benthic fossils are lacking. If the ostracods were benthic, then bottom conditions were probably dysoxic because ostracods carmot survive in oxygen-free environments

(Lethiers and Whatley 1994). Articulated ostracod carapaces from the Scs subfacies overlapped in cross-section, indicating benthic forms (Lethiers and Whatley 1994). Thus, rocks in the Scs facies that contained ostracods probably represent dysoxic conditions.

Hogler (1992a) interpreted her Biofacies 2 beds (comparable to the Sbl and Scs subfacies here) as representing dysoxic environments. However, one bed within the Scs subfacies 102

contained no ostracods or other visible macrofossils; that bed could indicate anoxic

benthic conditions.

In summary, sediments of the shaly fossiliferous limestone facies represent outer

shelf environments at just below storm wave base and deeper. Benthic oxygen levels

decreased through time and are reflected in the transition from bioturbated limestones

with abundant parautochthonous benthic fossils to non-bioturbated, calcareous shale and

limestone with few benthic fossils.

Pel-Bioclastic Limestone Facies (P*)

Rocks of the pel-bioclastic limestone facies include wackestones and packstones

with an abundant peloidal or pelleted fraction. Sediments of this facies contain less

terrigenous material than those of the shaly fossiliferous limestone facies. Two subfacies

are included in the pel-bioclastic limestone facies: peloidal bioclastic limestone (Pel) and

pelleted bioclastic limestone (Pbl). Ichthyosaur bones were not observed in the P facies.

Description

Peloidal bioclastic limestone subfacies (Pel. Figure 2.10). Medium-bedded black

micrites, wackestones, and rare packstones comprise the peloidal bioclastic limestone subfacies. These limestones weather blue-gray, brown, gray, or green (where altered by

intrusives) and are dark gray to black on fresh surfaces. Beds often occur in sets with a sharp, irregular base followed by coarse bioclasts (including whole fossils), swaley laminations, and a lighter-colored, grainy bioclastic top. Very thin weathered-out beds of 103

fossil hash are abundant in float near the tops of the beds. Some beds are amalgamated

and contain alternating fine and coarse-grained centimeter-scale bedsets.

Grains are medium- to poorly sorted and include peloids, bioclastic hash, rare

oolites, and rarely, whole fossils. Peloids are 0.2 mm long on average, sub-rounded, and

variably shaped. Cephalopods and halobiid bivalves, though uncommon, are the

predominant macrofossils and are usually preserved as impressions, recrystallized single

shells, or as mud and spar-filled shells, in the case of cephalopods. Echinoid fragments

and ostracods are occasionally visible in thin section.

Individual beds often fine upward. Where present, the basal, coarse part of a

fining-upward sequence tends to be poorly sorted. Smaller size fractions higher in the sequence each tend to be moderately well-sorted with densely packed, uniformly distributed bioclasts. In one slab, imbricated cephalopods were found in the coarser, basal

part of a fining-upward sequence, while halobiid bivalves were found in the middle part of the same sequence, oriented parallel to bedding.

In the upper exposures of this subfacies, peloids, oolites, and micritized grains increase in abundance. Pressure solution seams, stylolites, and calcite veins are common.

Some features that appear to be laminations on weathered surfaces are actually swarms of pressure solution seams oriented parallel to bedding. Other laminations are primary.

Pelleted bioclastic limestone subfacies (Pbl. Figure 2.11). Lithologies represented by the Pbl subfacies include cliff-forming, thick- and wavy-bedded micrite and wackestone. These rocks weather light blue or gray and are black to gray on fresh 104

surfaces. Well-sorted and uniformly shaped peloids (here termed pellets) and bioclastic hash characterize these rocks. Pellets are fine-grained, ovoid, and range in size from 0.08

- 0.4 mm, although most are around 0.1 mm. They are randomly oriented and do not appear compacted. Most of the bioclastic hash consists of disarticulated brachiopod shells and shell fragments, with some original microstructure preserved. Minor amounts of echinoderm and gastropod debris are present in some beds. Bioclasts form dense to loosely packed patches that are poorly sorted and have variable orientations. A few beds contain rare intraclasts with micritized fossils. Pressure solution seams, stylolites, and calcite veins are common. Some beds have undergone extensive dissolution based on numerous swarms of pressure solution seams.

Interpretation

The term "peloidal" is used in the loose sense when referring to the Pel subfacies, implying various origins of the peloids (Tucker and Wright 1990). Peloids of the Pel subfacies are irregularly shaped and occur with rare micritized bioclasts and ooids. Pel subfacies peloids probably originated via micritization of a variety of grains, including bioclasts, ooids, and fecal pellets. In contrast, the pelleted bioclastic limestone subfacies

(Pbl) contains unrimmed, ovoid grains of uniform size and shape that are probably fecal pellets (Fliigel 1982; Tucker and Wright 1990).

The Pel subfacies represents shelf environments between fair weather and storm wave base. Imbricated ammonoids, sharp bases, fining-upward sequences, swaley laminations, and amalgamated sequences indicate deposition by storm events. These 105

beds likely represent medial tempestites (Brett and Baird 1986), as some mud is present

and this subfacies is interbedded with Sbl and Scs subfacies representative of more distal

outer shelf environments. Peloids, micritized grains, and ooids were most likely

transported from more nearshore environments and deposited with fossils associated with

deeper water, such as ammonoids and halobiid bivalves.

The Pbl subfacies contains pellets of probable fecal origin that underwent early

lithification, based on their uncompacted appearance and random orientation. The

disarticulated and fragmented nature of bioclasts suggests deposition in or transportation

from an energetic environment. Beds of this subfacies are not condensed, because the

pelleted matrix, though cemented early on, is not bored or burrowed, and because one

kind of bioclast (brachiopods) predominates. Additionally, bioclasts do not appear to be

encrusted or bored. Intraclasts and rare bioclasts with light brown mud fill suggest

transportation from a different environment (Kidwell 1991).

Oolitic Bioclastic Limestone Facies (O. Figure 2.12)

Description

Rocks of the O facies include packstones and grainstones that form massive to

thick- and wavy-bedded cliffs. Beds weather gray to dark-gray and are dark-gray to

black on fresh surfaces. Lower occurrences of the O facies contain ooids almost exclusively, while higher occurrences contain a more diverse array of bioclasts. The O

facies culminates in a bioclastic echinoderm grainstone that grades into a small ledge of

brachiopod packstone. The grainstone is 9 meters thick and has faint, small- to medium- 106

scale cross-beds. The brachiopod packstone ledge contains small brachiopods in all

states of preservation but with most shells abraded. Some microstructure is preserved.

In the O facies, oolites generally contain only one lamina. Ooid nuclei consist of

peloids, micritized grains, and microfossils such as foraminifera. Brachiopods and

echinoderm fragments, especially crinoid columnals, are abundant in upper occurrences

of this facies. Other fauna include pelecypods, foraminifera, rare gastropods, and other

bioclasts. Preservation styles include recrystallization, internal molds, and micritic

envelopes. Common diagenetic features include syntaxial cements and pressure solution

seams.

Interpretation

The oolitic bioclastic limestone facies represents a shallow shelf environment containing carbonate shoals. Most lithologies, especially the grainstone units, contain

little to no mud, indicating current activity sufficient to winnow away very fine-grained

material. A preponderance of small, moderately sorted bioclasts also indicates current activity. Oolites indicate well-washed, shallow marine environments. The brachiopod ledge at the top of the grainstone likely represents colonization of a gravelly substrate.

Fossiliferous Limestone Facies (F. Figure 2.13)

Description

Massive to thick-bedded, cliff-forming, dark-gray to black whole-fossil wackestones and packstones occur in the F facies. Fossils are mud-supported. Burrow mottles are common, and burrow networks {IThalassinoides) are visible on the soles of 107

some beds. Large ostreid bivalves (Lopha cordilleram McRoberts 1998), large, high-

spired gastropods, large brachiopods, and echinoderm fragments are the most common

macrofossils of this facies. Ostracods occur rarely. Moderately to poorly sorted patches

of articulated shells, disarticulated shells, and fossil fragments occur in variable

orientations. Occasional thin (< 1 cm), crude laminations of bioclasts, some with convex-

up orientations, are visible. Fossils are recrystallized, although some ostreid and

brachiopod shell microstructure is preserved. Encrusters are rare, and borings lacking.

Pressure solution seams and microstylolites are common.

Interpretation

The F facies represents deposition in shelf environments in a quiet-water setting,

landward of carbonate ooid shoals but still flilly marine. The large size of fossils in this

facies compared to others in the stratigraphic section suggests stable conditions favorable for growth for extended periods of time. Similar, though not strictly analogous, quiet- water environments are observed today on the iimer shelf margin of the South Florida

Shelf (a rimmed, clastic-free shelf), where organisms are major contributors to the sediment budget (Sellwood 1986). Back-shoal environments were suggested for the

Middle Jurassic of north-western Europe (an epeiric sea), which includes beds with an analogous fauna to what is observed in the F facies (Sellwood 1986).

In summary, lithofacies of the marine part of the Luning Formation exposed in

West Union Canyon represent environments below storm wave base (S), between storm and fair-weather wave base (P), and above fair-weather wave-base (O, F). 108

Geochemical Results

TOC values ranged from 0.01 weight % to 0.85 weight %, with a mean value of

0.28 weight %. The sample with the highest TOC (field number 72995-1) gave Rock-

Eval pyrolysis results of S 1=0, S2=0, and S3=0.65 mg C02/g of rock sample (Figures

2.14 and 2.15, Table 2.5, Table B.l). Samples from the Scs subfacies had the highest average TOC contents, while samples from the Pbl and O facies have the lowest average

TOC contents (Table 2.5).

Facies trends for major and minor elements were similar to those for TOC

(Figures 2.14 and 2.15, Table 2.5, Table B.2). Samples from the Scs subfacies had the highest average major and minor element concentrations, while the P, O, and F facies contained the lowest concentrations. Ba, Cd, V, Zn, Na, K, and A1 in particular occurred in highest concentrations in the Scs subfacies. Of the 37 samples analyzed, 6 contained <

30 weight % Ca (Table 8.2). Five of the six samples were from the Scs subfacies. These five samples also had the highest TOC contents of all rocks analyzed. The sixth sample with < 30 weight % Ca was a bivalve internal mold from the Sla subfacies. Phosphorus values ranged from <0.1 to 0.12 weight % with a mean value of 0.06 weight % and highest concentrations were in samples from the Sla and Scs subfacies. Two bivalve internal molds from the Sla subfacies were not enriched in phosphorus or any other elements (Table B.2).

Correlation coefficients for the elements listed in Table 2.5 are given in Table 2.6.

I did not include results for Ca, Be, Co, As, Ag, Cd, Sn, Sb, W, Pb, Bi, Mo, or Ti in the 109

statistical analyses of geochemical trends. Most of the samples contained concentrations

of these elements that were either above (for Ca) or below (other elements listed above)

the detection limits of the chemical analysis (Table B.2). The correlation coefficients and

average elemental concentrations of each of the subfacies will be compared with

elemental concentrations from other marine environments in the discussion.

Paleontological Results

Below, I describe major features of the invertebrate fauna in terms of the

lithofacies discussed above. These features are also summarized in Figure 2.16, which

used abundance data from the bulk macrofaunal analysis of Hogler (1992a, see that paper

for details on bulk analysis samples and trophic and biofacies assignments). My field

observations agreed with the general conclusions of Hogler's (1992a) analysis. Her fossil

invertebrate abundance data were collected from exposed beds in the Luning Formation

in West Union Canyon. Despite the sampling bias to exposed rock, these data are useful

for describing the general faunal trends (Figure 2.16).

In general, suspension feeders dominate the preserved fauna, and suspension-

feeding bivalve molluscs are the most common assemblage constituents. Exceptions

occur in the O and Pbl facies, where brachiopods dominate (Figure 2.16). Gastropods

and epifaunal and shallow-infaunal suspension-feeding bivalves dominate the Sla

subfacies. In the deeper-water Sbl subfacies, ammonoids and deposit-feeding bivalves are important components of the fossil assemblage. Infaunal benthic fossils are still rarer

in the Scs subfacies, where anmionoids, halobiid bivalves, and/or ostracods are the main 110

(sometimes only) components of the assemblage. The Pel subfacies contains both

autochthonous ammonoids and halobiid bivalves and allochthonous echinoderm

fragments. The O and Pbl subfacies are dominated by brachiopods and crinoid fragments

(O facies). The F facies is dominated by ostreid bivalves, brachiopods, and gastropods.

Corals, sponges, and foraminifera are rare in all facies.

Ichthyosaur fossils are most abundant and well-preserved in the Sbl subfacies. In order of decreasing abundance, bones are also found in the Sla and Scs subfacies. I did

not observe bones in other lithofacies, although one Scs/Sbl occurrence (field number

73095-10) is located in the upper 1/3 of the stratigraphic section. Hogler (1992b) reported bone occurrences throughout the marine part of the section, but bones were most abundant and well-preserved in her Biofacies 2 (Sbl/Scs subfacies of this dissertation). Ill

DISCUSSION

Environments Of Deposition

Sedimentological Aspects

Rocks of the marine part of the Luning Formation in West Union Canyon reflect a

range of environments, from deeper shelf muds to carbonate ooid shoals (Figure 2.17).

There is no evidence for shelf-break or slope deposits at BISP. Beds are not slumped and

soft-sediment deformation is lacking. No resedimented carbonates are present.

Transported carbonate rocks are present, though, farther west (across structural

boundaries) in the Lovelock and Sand Springs lithotectonic assemblages (Oldow, 1984,

Paradise subterrane of Silberling et al. 1987). These rocks could represent the slope and

basinal deposits off of the Luning shelf (Oldow et al. 1993). The presence of resedimented carbonates west of BISP supports the shelf interpretation for Luning rocks.

The Luning Formation at BISP reflects a range of energy conditions, from below storm wave base to above fair-weather wave base. The lower half of the composite stratigraphic section records environments below or just at storm wave base, reflected by the lack of primary sedimentary structures, the presence of whole bivalves, and the lack of a preferred orientation of fossils in the Sla, Sbl, and Scs subfacies. The upper half of the composite section records shallowing from below storm wave base to above fair- weather wave base, reflected by the presence of swalely laminations, peloids, and bioclasts (as opposed to whole fossils) in the Pel subfacies and poorly formed ooids in the 112

O facies. The lower two-thirds of the section was formed in middle-outer shelf environments, while the upper third was formed in middle-irmer shelf envirormients.

No facies in the section contain features that indicate lag surfaces or condensation beds. Calculated sedimentation rates were highest for S facies sediments and none of the calculated rates were exceptionally low (Table 2.2). Notably, the beds with the most abundant ichthyosaur bones (Sbl subfacies) do not have the lowest sedimentation rates, arguing against accumulation of skeletons by condensation. The calculated rates are typical of ancient shallow-marine carbonate environments (see Table 1-1 in Wilson

1975). Carbonate forms today at rates of this magnitude and higher in temperate and tropical pelagic, shallow subtidal, and open shelf areas (Rao 1996). The rates for the

Luning Formation at BISP are minimum estimates because solution seams and stylolites are visible in many samples throughout the section and compaction has not been taken into account. These rates are comparable to the highest sedimentation rates estimated for the Lower Oxford Clay of Callovian age in England (Hudson and Martill 1991; their estimates ranged from 0.09 - 1.5 cm/1000 yr, assuming 430,000 years/ammonoid subzone), a rock unit that was deposited on a flooded shelf and is thought to have experienced elevated productivity related to terrigenous nutrient flux.

Calcareous shales (Scs subfacies) alternate with peloidal, bioclastic carbonates

(Pbl subfacies) containing rare ooids (probably transported from nearshore envirormients) in the 300-450 meter interval of the composite stratigraphic section (Figure 2.5). This facies interfingering presents an interesting situation. The clastic material in the shales 113

was presumably terrestrial in origin, whereas the ooids presumably represent a current-

washed setting relatively free of clastic material. Clastic transport appears to have

decreased upsection, based on the silty appearance of Sla and Sbl rocks versus the

relatively clean appearance of P and O rocks in hand sample and thin section. Major

element concentrations associated with clastic material also decrease upsection (see

below). Development of an ooid shoal, represented by the O facies, likely occurred as sea level rose and more clastic material became trapped in river mouths and estuaries.

Evidence of oolites is only found in the P and O facies, and no facies lower in the section

contain transported oolites. The ooid shoal was probably a local, somewhat short-lived feature because of the evidence for abundant clastic transport at contemporaneous

Luning Formation localities (e.g., the Pilot Mountains).

Geochemical Aspects

The distribution of major elements in the section suggests that terrigenous silt input decreased or was diluted by other components over time. Concentrations of A1 and

Ti, which estimate the detrital content (Sugisaki 1984; Brumsack 1986; Piper and Isaacs

1995), were higher in the S facies and in the Scs subfacies in particular, and lower in facies farther upsection (Figures 2.14 and 2.15, Table B.2).

Samples from the Scs subfacies had the highest TOC contents, consistent with its fine-grained nature (Boggs 1992; Tyson 1995). Three of these samples contained >0.50 weight % organic carbon, which can be considered organic-rich for carbonate rocks

(Parrish 1995), although below the average organic content of shales (1.0 weight %, 114

Boggs 1992). However, the Rock-Eval pyrolysis results of no SI or S2 and higher S3

indicate highly mature, oxidized organic matter with poor hydrocarbon-generating

potential. This pattern of Rock-Eval results can occur with weathered outcrop samples

(Peters 1986), with the presence of refractory organic matter such as remains of vascular

plants, or with thermal alteration of sediments (Tissot and Welte 1984). Conodonts from

ammonoid-bearing rocks in the section showed an alteration index of 5 (N.J. Silberling,

personal communication 1996), which indicates that the rocks experienced temperatures

between 300 and 400° C (Epstein et al. 1977). Metamorphic mineral assemblages in

nearby Early Triassic - Late Permian volcanic rocks (N.J. Silberling, personal

communication 1996) also indicate fairly high temperatures during diagenesis. High

temperatures could have driven out much of the organic carbon in the rock, meaning that

the TOC results presented here are minimum estimates of the original organic carbon

content of the rock. On a regional scale, diagenetic conditions in the MMP were at one

time favorable for petroleum generation, as liquid hydrocarbons are found in ammonoid

chambers from the late Anisian Fossil Hill Member of the Favret Formation (Nichols and

Silberling 1977). This formation represents a deep shelf envirorunent similar to that of the Luning Scs subfacies, and outcrops today in the Augusta Mountains northwest of the

Shoshone Mountains.

Minor element concentrations can indicate benthic redox conditions (Calvert and

Pedersen 1993). High Mn concentrations coupled with low V, Cr, Ni, Cu, Zn, Mo, and

Cd concentrations suggest oxic bottom water (Calvert and Pedersen 1993). At BISP, Mn 115

is slightly higher in the Sla subfacies and suggests oxic benthic conditions. Mo, Cd, Cu, and Zn are higher in the Scs subfacies, while Mn is low (Tables 2.4, B.2), suggesting anoxic sediments and/or bottom waters during deposition.

The correlation coefficients calculated for 18 samples of 19 elements (Table 2.6) are largest for TOC/Zn, Cr/Cu, Ni/Cu, Al/Sc, Sc/La, and Cr/Fe. The first 3 element pairs are most likely related to biogenic marine sources, whereas the Al, Sc, and La pairs are most likely related to detrital sources. Table 2.6 also contains some fairly high coefficients between one element each of detrital and biogenic-marine sources (e.g.,

Al/Cu). These correlations could reflect the detrital minor element contribution

(Medrano and Piper 1992; Piper 1994) or the effects of diagenesis and weathering.

Several studies have utilized a normative-calculation procedure to partition minor elements into major mineral components (Medrano and Piper 1992; Piper 1994; Piper and Medrano 1994; Piper and Isaacs 1995), but Luning data are too few for those calculations. In general, Luning samples do not show high elemental concentrations except for certain elements in the Scs (Fe, Zn) and Sla (Fe, Mn) subfacies (Tables 2.4,

B.2). Zinc is associated with sedimentation of organic carbon (Piper 1994) and Mn is associated with benthic oxygen conditions (Calvert and Pedersen 1993). Iron is informative of benthic oxygen levels if the relative proportion of oxidized to reduced forms is known (Raiswell et al. 1988; not measured in the analyses presented here).

Sampling constraints might have biased geochemical results. Sampling was conducted along the traverse used to measure the section and was limited to outcrops. 116

Trenching of shale horizons yielded samples that were still well weathered. Use of weathered samples means that geochemical analyses might not be representative of the rock unit sampled. Systematic bias was introduced by using small hand sample splits for analysis. This further increases the chances of analyzing non-representative portions of the rock unit. However, I was careful to choose the least weathered samples and to remove calcite veins from the sample chips. Average elemental concentrations for lithofacies compare favorably with reference material compositions of similar lithology

(Table 2.7).

Paleontological Aspects

Whereas the taxonomic composition of the biota changes over the stratigraphic section, suspension feeding is the dominant strategy in the lower and upper thirds of the section (Sla subfacies; P, O, and F facies). Camivores and/or scavengers (represented by ammonoids) dominate the Sbl and Scs subfacies, with some suspension feeders

(halobiids) present (Figure 2.16). Throughout the section, most faunal elements are autochthonous or parautochthonous except for bioclasts in the P and O facies. Whole fossils (namely brachiopods) that are present in these facies are in place or have been transported very little, particularly in the O facies.

Ichthyosaur remains are most common in the Sbl subfacies; skeletons are partially articulated and bones are somewhat abraded, suggesting some, but not prolonged, exposure on the seafloor. There is no evidence for condensation or bone lags. Rather, bone deposition occurred throughout Sbl and to a lesser extent, Sla sedimentation. As 117

skeletons are often partial, rather than complete, and skulls are relatively rare compared

with ribs and limbs (Camp 1980), skeletons could have been deposited in the BISP area

after having been refloated and current-drifted, as Schafer (1972) observed for whales in

the North Sea. The abundance of skeletons and their occurrence over several horizons

indicates that large ichthyosaurs inhabited the region over a prolonged period of time

(Hogler 1992b). Ichthyosaur bone absence in the P, O, and F facies could reflect a shift

in ichthyosaur distribution related to development of favorable habitats elsewhere in the

region. During deposition of the shallower Luning sediments in the upper part of the

section at BISP, deeper-water shelf sediments similar to the Sla and Sbl subfacies were

being deposited in the Luning now exposed in the Pilot Mountains, and ichthyosaur

remains are common there.

Upwelling Characteristics

Upwelling is a complex process with many potential effects on the sedimentary

record (see Chapter 1). Presence of the macrofaunal characteristics listed in Table 2.1

can indicate upwelling in deposits that lack geochemical criteria. Several aspects of

Luning Formation sediments at BISP suggest that upwelling could have influenced their

deposition. In the following sections I discuss the Luning Formation in terms of cool-

water conditions, oxygen-minimum-zone edge effects, and eutrophic conditions.

Cool-Water Conditions

The lack of corals and calcareous algae, especially in shallower facies of the BISP section, suggests cool water, somewhat of an anomaly considering the subtropical 118

paleolatitude suggested for this area in the Late Triassic (paleolatitudes based on faunas

Tozer 1982). If upwelling operated in the MMP basin, then one would expect cooler water faunas and sediments. Thin coral and sponge buildups are present in the lower member of the Luning Formation at the Pilot Mountains (Stanley 1979). Stanley (1979) suggested that the organisms forming the buildups were ahermatypic, enabling the animals to cross Panthalassa and inhabit deeper and/or more turbid environments.

Calcareous algae are also absent from this locality (in contrast to Tethyan localities), indicating cool and/or deep water (Stanley 1979; 1988a). On an even larger scale, cooler waters and/or increased turbidity associated with tectonic activity could explain the limited the growth of western North American corals as compared with the large reefs developed in contemporaneous Tethyan localities (Stanley 1988b).

No taxa from BISP exhibit clear cold-water affinities, though several are cosmopolitan in distribution. On the other hand, to my knowledge no taxa from BISP are known to be limited to low latitudes. Late Triassic Halobia sp. are found in localities with polar to tropical paleolatitudes (McRoberts 1997). The Late Triassic oyster

Gryphaea is known from high and middle paleolatitudes and from low paleolatitudes in western North America but not in Tethys (McRoberts 1992). This oyster is found in both the Pilot Mountains (McRoberts 1992) and at BISP (McRoberts, personal communication

1998). McRoberts (1992) suggested that Gryphaea inhabited cool and/or deep water, based on the associated faunas in the Luning Formation, the Shublik Formation (North

Slope, AK), and the Martin Bridge Formation (Wallowa Mountains, OR). 119

The presence of ooids in the O facies and to a lesser extent the Pel subfacies

indicates warm-water conditions. However, the ooids are relatively poorly developed

(Figures 2.10 and 2.12) and occur in thin intervals compared with the other facies (Figure

A.5). A mix of warm- and cool-water indicators does not preclude upwelling. Brandley

and Krause (1997) found faunal and sedimentological indicators of warm-, cool-, and

cold-water on a Carboniferous, equatorial carbonate ramp and concluded that upwelling

resulted in the cool- and cold-water facies while solar heating of nearshore environments

resulted in the warm-water facies.

Edge Effects

Rocks of the Scs subfacies had the highest TOC values and contained the fewest

benthic macrofauna of all the subfacies. Sediments of the Scs subfacies were thus

deposited under the lowest benthic oxygen levels of any rocks in the section. In the

Luning sediments at BISP, the presence of the upper edge of an oxygen-minimum zone

(OMZ) is reflected by a decrease in preserved benthic organisms and an increase in

organic carbon content in the Sbl and Scs subfacies. Faunal density is greatest in mid-

shelf facies (Sla) that were presumably at or just above the upper edge of the OMZ

(Figure 2.16), similar to the density pattern of major taxa with depth from today's central

California OMZ (Vercoutere et al. 1987). Other marine localities in the Luning

Formation and in the overlying Early Jurassic Gabbs-Sunrise Formation have similar

faunal density patterns and facies (Laws 1982), suggesting that the OMZ was laterally and temporally persistent in the basin. 120

None of the rocks observed at BISP were barren of fossils. Samples from the Scs facies often contained only ostracods that appear to be benthic, indicating dysoxic conditions (Lethiers and Whatley 1994). This observation also supports the interpretation of the Scs facies representing the upper edge of an OMZ (as opposed to the whole OMZ, including the very-low-oxygen core). Hogler (1992a) came to the same conclusion of a dysoxic but not anoxic environment for her biofacies 2 (Sbl/Scs subfacies), though she used an epibenthic habit for Halobia sp. rather than the presence of ostracods to support this conclusion.

Eutrophic Conditions

The dominance of suspension-feeding animals over most of the Luning section indicates an adequate supply of food particles suspended in the water column (Figure

2.16). This dominance, coupled with a lack of zooxanthellate corals, can indicate high productivity from either upwelling or nutrient-rich runoff (Hallock and Schlager 1986;

Brasier 1995). Beds with abundant recognizable fecal pellets (Pbl subfacies) indicate biogenic input to the sediment that exceeded organic recycling (Porter and Robbins

1981). Other facies with clotted fabrics (S and F facies) might also contain fecal pellets compacted beyond recognition.

Ammonoids at BISP show morphologies typical of nektonic and planktonic habits

(Figure 2.18) (Westermarm 1996). A mixture of ammonoid ecologies is present in the

Sla subfacies, including nektonic and vertical planktic migrant forms. The proportion of vertical planktic migrants increases in the Sbl subfacies. Lower beds of the Sbl subfacies 121

are dominated by a single genus of vertical migrants, Tropites sp. In upper occurrences

of the Sbl subfacies, several taxa of vertical migrants are present, and one taxon with a

highly evolute shell, Tropiceltites columbianus, was possibly a planktic drifter that spent

its life in the epipelagic environment. The dominance of possibly planktic forms in the S

facies suggests a plentiful epipelagic food supply.

The geochemical composition of rocks often varies widely (Fairchild et al. 1988).

Rock units regarded as upwelling deposits are no exceptions. For example, phosphate

values across facies of the Permian Phosphoria Formation vary from 0.16-39 weight

percent (Medrano and Piper 1992). Luning samples had low P contents (0.12 weight

percent maximum. Table B.2). However, P alone is not a sure indicator of an upwelling environment because phosphate is found in abundance in only 2 of the 5 major coastal

upwelling areas today (see Chapter 1). Single samples of argillaceous limestone from the

Phosphoria Formation and calcareous mudstone from the Monterey Formation have compositions similar to that of the Scs subfacies (Table 2.7, rows 3,15, 16). V, Cr, Ni, and Zn compositions are higher in the Phosphoria and Monterey samples, but the acid dissolution technique used on the Luning samples might not have extracted all of the V and Cr present.

The marine fraction of several minor elements, including Cd, Cu, Ni, Zn, V, Cr, and Mo, either have a nutrient-type profile, suggesting that they taken up extensively by plankton, or, in the case of Mo, are used by plankton but not in quantities that deplete the element's concentration in surface waters (Piper 1994). Piper (1994) suggested that 122

plankton contribute most of the marine minor-element fraction, even in ancient sediments with high metal concentrations. Calvert and Price (1983) recognized an association of

Mo, Cu, Ni, S, and C in Walvis Bay, where persistent upwelling occurs. While Luning

Mo and Cd values were not included in the statistical analysis, Cd levels in some samples of the Sbl, Scs, and Pel subfacies exceeded those of average shale and were comparable with 2 samples from the Monterey Formation (Table 2.7). Similarly, Mo levels in the

Sbl, Scs, O, and F subfacies exceeded levels in average shale and approached levels in 2 samples from the Phosphoria Formation (Table 2.7). Of the 7 Luning subfacies, the Scs subfacies had the highest levels of Zn, which was also highly correlated with TOC content (Figure 2.5).

Synthesis

In summary, lithologic evidence at BISP indicates shelf deposition in cool, deep, and/or turbid environments. A depauperate benthic fauna and the presence of ostracods in the Scs subfacies suggest that the OMZ was near this area at the time of Scs subfacies deposition. Modest organic carbon enrichment in the Scs subfacies and association of

TOC with Zn suggests moderate productivity and biogenic marine sources of some minor elements. Paleontological results indicate presence of cosmopolitan forms found in low to high latitudes, but no taxa restricted to low latitudes. Fecal pellets and clotted fabrics suggest high productivity at least during some depositional intervals.

Given the above, several possible scenarios exist for Luning deposition (Figure

2.19). The scenarios include (1) sedimentation without high productivity but with a well- 123

developed OMZ, (2) sedimentation with high productivity influenced by upwelling, and

(3) sedimentation with high productivity influenced by runoff.

Scenario 1

In scenario 1 (Figure 2.19, Table 2.8), productivity would reach average marine shelf levels but would not be enhanced by upwelling or runoff. The OMZ would result from stagnation of the basin and waters below the OMZ likely would be devoid of benthos. The most oxygen-depleted facies would be expected in the central, deepest part of the basin (Parrish 1995). In scenario 1, the abundance of ichthyosaurs could result from concentration of bones on the seafloor via slow sedimentation and formation of lag deposits (Norris 1986). Alternatively, many bones could be preserved by rapid burial via high influx of terrigenous sediment, but then the reasons for having a large supply of bones in an area without much food would need to be explained. Terrigenous influx would not be a prominent agent for burial during P and O facies deposition, because the ooid shoal presumably existed at that time.

Taylor et al. (1983) suggested that an expanded OMZ contributed to depauperate, offshore faunal associations found in some members of the Triassic-Jurassic Volcano

Peak Group (Gabbs and Sunrise Formations) that overlie the Luning. However, depauperate benthic faimas can also form under OMZ's associated with upwelling zones.

Additionally, offshore deposits in the Volcano Peak and in the Luning still represent shelf environments and are thus offset from the center of the basin, where one would expect the most oxygen-depleted conditions under scenario 1. Contemporaneous rocks ("the 124

mudpile") now exposed west of BISP across structural contacts likely were deposited

under hemipelagic conditions in a rapidly subsiding basin (Oldow et al. 1993). Benthic

marine fossils are sometimes rare in the mudpile (Speed 1978a), but the huge volume of

sediment deposited into the basin suggests rapid deposition rather than low oxygen as the

main factor influencing abundance. While the condition of ichthyosaur remains at BISP

indicates some exposure, the remains are heterogeneously distributed in both

discontinuous limestone beds and calcareous shales (Sbl subfacies, see also Camp 1980).

Evidence for condensation and lags are lacking.

Scenario 2

In scenario 2 (Figure 2.19, Table 2.8), productivity enhanced by upwelling would

result in an intensified OMZ, with oxygenated waters below the OMZ. Depending on

basin geometry and current dynamics, a range of sedimentary environments could result.

With a shallow shelf, the situation off of northwest Afnca today, organic matter is concentrated on slope and basin deposits, whereas coarse material predominates on the shelf. In contrast, off of southwest Afiica, upwelling and deposition of biogenic sediment occurs along the inner shelf (Summerhayes 1983). Elevated productivity would likely create sediments high in organic matter, P, and minor elements associated with plankton

(e.g, Cu, Ni Zn, Piper 1994). Upwelling of cooler, nutrient-rich water would permit cool-water taxa to live in low-latitude regions. Arctic brachiopods are present in the

Permian Phosphoria Formation, a rock unit deposited m the equatorial belt and interpreted as an ancient upwelling zone (Wardlaw and CoIIinson 1986). 125

Data from the Luning Formation are consistent with Scenario 2 for a moderately

productive coastal upwelling system. Higher TOC and a depauperate benthos in the Scs

subfacies suggest the presence of an OMZ on the distal part of the shelf. BISP facies, all

shelfal, contain more mud than coarse-grained material, so current winnowing on the

mid-outer shelf was probably minimal, unlike the situation today off of northwest Africa.

Rocks of the Scs subfacies at BISP contain, at a minimum, modest enrichments of organic carbon and an association between Zn and TOC, suggesting a marine origin for at least some organic carbon. Enrichments of other elements are lacking, but it is clear from conodont alteration, metamorphic minerals, the overmaturity of TOC, and the abundance of solution features that rocks at the BISP have undergone fairly extensive diagenesis, possibly altering their inorganic composition.

Macro fossils from the Late Triassic Shublik Formation, deposited at high paleolatitudes, do not include exclusively cold water forms but do include the cosmopolitan Halobia sp. (Parrish 1987a). Faunas from BISP also include cosmopolitan

{Halobia sp., Palaeocardiata sp.l) and possibly cool-water {Gryphaea sp.) forms and a lack of warm-water calcareous algae. Macrofauna are abundant over most of the section and suspension-feeders dominate, suggesting a plentiful food supply.

Overall, sedimentological and paleontological evidence favors scenario 2 over scenario 1. However, the lack of geochemical enrichment of P and/or other marine- source minor elements suggests that upwelling, if present, was not intense. The 126

overmature nature of the TOC at BISP could also result from heating associated with diagenesis or a high proportion of land-based, refractory organic matter.

Scenario 3

In scenario 3 (Figure 2.19, Table 2.8), high productivity would occur in the basin but would be driven mainly by terrigneous input from river systems. The positive effects of runoff generally occur seaward of the river mouth. Runoff can negatively affect benthic production by erosion of the substrate, smothering, salinity reduction, and anoxia caused by water stratification (Alongi 1990). In the East China Sea off of Changjiang,

Rhoads and others (1985) found that benthic communities stabilized and production increased seaward of the river mouth, particularly in the mid-shelf region. Closer to the river mouth, high, rapid sedimentation smothered benthic communities fairly often, overriding any nutrient benefits (Rhoads et al. 1985). A similar situation exists for the benthic communities near the Amazon River (Aller and Aller 1986).

Runoff (particularly if rich in dissolved nutrients in solution) can enhance productivity if the water column is adequately stable and transparent (though runoff can be nutrient-poor in tropical areas, Raymont 1980). Today in the Banda Sea, seasonally alternating upwelling and runoff associated with a monsoonal climate create a depositional situation where runoff raises the sedimentation rate and helps preserve biogenic sediments (van Waveren and Visscher 1994). Martill and others (1994) proposed that the Jurassic Oxford Clay, an organic-rich, fine-grained unit with abundant micro-, macro-, and mega-faunas, experienced high productivity associated with runoff. 127

Terrestrial and/or nearshore organic matter can make a significant contribution to marine sediments (Hedges 1992).

In the Triassic, the supercontinent Pangea was distributed symmetrically about the equator and was ideally situated for creating intense monsoonal circulation (Parrish

1993). East of the MMP, the Late Triassic Chinle Formation contains lithofacies and floras that suggest seasonality (Dubiel et al. 1991). With a monsoonal climate, it is possible that seasonal terrigenous input boosted productivity and/or enhanced preservation of upwelling sediments generated during the dry season, as in the Banda Sea today (van Waveren and Visscher 1994). Many of the same productivity indicators would be expected for scenarios 2 and 3. Potentially more terrigenous debris would be present in scenario 3.

Fine-grained terrigenous debris is abundant in the S facies at BISP. Large volumes of fine-grained clastic sediment were deposited at localities west of BISP in hemipelagic facies of the west-central MMP (Speed 1978a). Luning Formation localities each record a period of thick clastic deposition, indicating that fluvial systems drained into the basin (see "Lithology, Age, and Paleontology of the Luning Formation" above).

The potential for runoff-enhanced productivity thus remains for the Luning Formation, mainly for the lower half of the section (S facies). During P and O facies deposition, however, runoff was probably much lower as the carbonate ooid shoal was developed at this time. Runoff-enhanced productivity was also possible for the F facies, which was landward of the shoal. 128

The modest TOC enrichment, presence of the OMZ, and presence of possibly

cool-water taxa support scenarios 2 and 3 for the lower half of Luning Formation

deposition. Further data on clastic and organic matter composition would help delineate

between scenarios 2 and 3. Given the sedimentological and floral evidence on land for a

seasonal climate in the Late Triassic of western North America (Dubiel et al. 1991), it is

likely that terrigenous input and marine upwelling influenced productivity in the lower

half of the Luning Formation at BISP, if the ooid shoal did not develop until deposition

of the P facies. If the ooid shoal existed during the whole of Luning Formation

deposition, then runoff-enhanced productivity would be effective only for the F facies. In

that case, moderate-intensity coastal upwelling (scenario 2) would be the most likely contributor to eutrophic conditions in the Luning Formation at BISP. 129

CONCLUSIONS

In conclusion, sedimentological, paleontological, and geochemical evidence

support the idea of eutrophic conditions in the Luning Formation at BISP. While Luning

Formation rocks in the study area do not show "classical" upwelling facies (phosphorite,

glauconite, biosiliceous sediments), modest organic carbon enrichments were observed.

More importantly, several different kinds of macrofaunal evidence provide evidence of

elevated productivity, including a predominance of suspension feeders, dominance of

several cosmopolitan and possibly cool-water taxa, and dominance of planktic taxa in

facies representing deeper and more oxygen-poor envirorunents. Lack of coral buildups

and lack of calcareous algae also supports the interpretation of enhanced productivity.

Macrofaunal evidence can be used to interpret the relative influence of

productivity in the geologic record (Table 2.1), especially when geochemical data are

lacking or of poor quality. But macrofaunal data are more than just substitute upwelling

indicators. As shown in the introduction, macrofaunal responses to elevated productivity are sometimes the only preserved indicators of upwelling. Additionally, recognizing the faunal contribution to the upwelling record allows identification of a new class of

upwelling deposits—those where only the faunal response to productivity is preserved, as in the Pliocene Pinecrest Sand of Florida and the Northwest Afncan upwelling zone today.

Moderate-scale upwelling likely contributed to eutrophic conditions during

Luning deposition. Productivity in the lower half of the Luning Formation at BISP could 130

have been enhanced by clastic input also. However, for runoff to have made a significant contribution to productivity in the lower half of the Luning, the ooid shoal facies (O) must have been a short-lived feature. Further study of clastic and carbonate facies across localities can help determine the relative contribution of runoff versus upwelling to productivity of the Luning Formation at BISP and elsewhere.

Marine vertebrates are present in the proto-MMP in the Early Triassic (Massare and Callaway 1994), and are known in abundance from the Middle and Late Triassic

(Table 2.9, see also Leidy 1868; Nichols and Silberling 1977; Stanley 1979; Laws 1982;

Silberling and Nichols 1982; Sander et al. 1994; 1997). Reptile fossils increase in abundance southward with time, reflecting basin development and perhaps food availability. The present study shows that productivity could have affected sedimentation and faunas in the Luning Formation at BISP. As similar lithofacies and faunas are found in Middle and Late Triassic rocks of the eastern MMP, it is likely that these areas also experienced enhanced productivity. Accreted to cratonic North America (.Borderland Late Triassic (about 215 Ma) Stikine Backarc Basin CRATON MMP ilS Lunms

Marine Remnant Uplands AfG Coastal Plain

OCEAN Chime Fm

200 mi I 300 km

Fluvial paleocurrent trends

FIGURE 2.1. Tectonic setting of western North America in the Late Triassic, after Blakcy, 1997 (maps available at http;//vishnu. glg.nau.edu/rcb/tripaleo.html). The MMP was locatcd at about 15° north during the Late Triassic (Tozer, 1982). See also Lawton, 1994 Ul 120® W II8'W Fenccmaker Thmst Lithotectonic assemblages of Oldow et al., 1993 shown in Humboldt italics.

Locality abbreviations: Lovelock nCM northern Cedar Mountains sGVR southern Gabbs Valley Range nPM northern Pilot Mountains nPR northern Paradise Range sSM southern Shoshone Mountains

PI pre-Tertiaiy sedimentary Sand - 39°N Springs and volcanic rocks that include the Luning Fm. Luning Pine Nui Thrust • Granite Pamlico Lumng nCM 1 Thrust fault (dashed » where inferred)

^'^Sierra Nevada \ Strike-slip fault (dashed Batholith Gold Range where inferred)

SO km

FIGURE 2.2, Outcrop map of the southern Mesozoic marine province showing areas where the Luning Formation is exposed (as well as other pre-Tertiary sedimentary and volcanic rocks). Also shown are the lithotectonic assemblages of Oldow et al. 1993. The Pamlico and Luning assemblages roughly correspond to the Paradise subterrane ot Silberling et al. 1987. The whole MMP corresponds to Sonomia (Speed 1978b) See text for details and figure 2.1 for the western North American context. Modified from Oldow et al. 1993. E Paradise S Shoshone N Cedar N Pilot S Gabbs Dolomite, ± algal laminations Range Mountains Mountains Mountains Valley Range Limestone, ± bioclastic or fossiliferous Late Norian • RQLVP:iSsgySlif^,(!9!flida.0i 2of__QJdpwejjai ^ J 993^ Early Norian Conglomerate, sandstone, & 1 argillite (fine-grained rx often metamorphos^ to homfels) 5b ED Impure limestone & shale (± calcareous)

Unconformity 4a 5c The Luning Fm. in Major fault contact &=3 the S Gabbs Valley Range confomiably 8=3 Ichthyosaur bone 3a^ , 6=3 und^ies the Gabbs (Only Shoshone Mts. Fm., which contains occurrences have been surveyed 2a I icbthyosaur bone in detail-see Camp, 1980.) in micritesof the 4b lower member (mL Norian) (L.aws, Biostratigraphy 1982). (e=early, m=middle, l=late; E =

%*• •/. •* Early, L = Late) Outcrops arc • .* • .* • .• •• .* • .• extensively intnided • %. •,. •. and metamoiphosed. 1. conodonts, < L Camian No measured section '.\\\\\ Silberling and John, 1989 available. Thicknesses r.*' •/.\ 2. welleri Zone, m L Camian shown are for vVVv'.V Illustrative puiposes 2a. ammonoids, Silberling, 1959 only. 3. macrolobatus Zone, I L Camian 3a. ammonoids, Silberling, 1959 4. kerri Zone, e E Norian 4a. ammonoids, Silberling, 1959 rvm/ 4b. ammonoids. Brown. 1986 600 4c. ammonoids, Siliberling, 1984 Outcrops are highly 4c 5. magnus Zone, I E Norian 5b. molluscs. Brown, 1986 12 intnid^ and u metamoiphosed. 5c. ammonoids, Silberling, 1984 Bone occurence 2 from D. Montague- Judd, unpublished Outcrops are exposed in 13 complexly field notes. folded thrust nappes (Oldow, 1981) 0^ FIGURE 2.3. Chart showing lithologic variation among Luning Formation localities shown in figure 2.2. Thicknesses vary within localities Stratigraphic data from Laws. 1982; Oldow et al., 1993; Silberling and John, 1989; Speed et al., 1989; and Stanley, 1979. Upper Carbonate 7400 •

[I'Xj Calcareous Shale

Shaly Limestone .7600.

elastics

\ Major Faults

Measured stratigraphic section 1. Campground Section 2. Fossil Hill 3. Boundary Hill 4. Furnace Hill Visitor 5. Brachiopod Ridge

Nevada

c o Z

H Berlin-Ichthyosaur j ToiyabeNat'l. 0 feel 1000 State Park • • I Forest • Contour interval 200 feet FIGURE 2.4. Geologic map of the Luning Formation in West Union Canyon, Shoshone Mountains. Informal members of Silberling, 1959. Minor faults are not shown. Modified from Silberling, 1959. 135

Calcareous Ammonoids shale

Bivalves Bivalve laminations Lopha cordillerana v/V\^ McRoberts

Fossil Brachiopods hash

O Echinoderms Peloids • Crinoid columnals

00 * 1% Pellets Gastropods

® Ooids Ichthyosaur bone Ostracods Burrow mottling Wacke=wackestone Pack=packstone Grain=grainstone

FIGURE 2.5. a) Legend for composite, generalized stratigraphic column of the Luning Formation at West Union Canyon, Shoshone Mountains. b) (next page) Generalized stratigraphic column. Covered intervals are not shown. Petrographic observations such as peloids and ooids are shown. See text for lithofacies descriptions. Column includes ichthyosaur occurrence data from Camp (1980) and my field notes, and biostratigraphic information from Silberling (1959) and Silberling and Tozer (1968). Scale: 1 cm = 33.3 meters 136

FIGURE 2.5.b

500-- ••tr* •ir* mic*

400-

o M ^ • • • •

Guembeliies o zone .... lower lower 300- • —""" Norian Sbl o

Pel o /O r.Tstufl.-.-'. O 4^ ^ -"Oxv oa» 200- ^ O M

&=2 Klamathites ^ ^ J® ® i macrolobatus zone 100- Sbl upper upper Camian Scs ^ ^ ® J) Klamathites 1 schucherti Sla i2u zone u middle upper s f^o Camian T\ Facies I. Campground 2. Fossil Hill 3. Boundary Hill 4. Furnace Hill 5. Brachiopod Ridge 650 Ammonoid Zones _ a. Klamathites schucherti b. Klamathites macrohbatus c. Guembelites sp.

520-

390- •60694-30- •60794-1 -71195-32

•71195-28'

•71195-23 •60694-1

—71695-34- •60594-17 130- 71095-1

.0494-20 •60594-5

Cqnclation Line (Fig. 2.4) FIGURE 2.6.2 6 Chart showing correlation among stratigraphic sections in the marine part of the Luning Formation at West Union Canyon. See Appendix A for columnar sections and key to sample numbers. FIGURE 2.7. Representative samples of the Sla subfacies; a) 60494-8, wackestone ledge interbedded with well-weathered argillite, b) 70895- 3.91, Palaeocardiala sp. packstone with whole bivalves weathering out of muddy limestone, (hammer head 18.5 cm long), c) 70895-3.91, acetate peel. Note whole, spar-filled bivalve in lower right, disarticulated and fragmented shells, round echinoderm debris (upper right), and fine-grained matrix, d) 70895-3.94, packstone, thin section. Note small gastropod with darker, finer-grained fill, fragment of recrystallized bivalve shell, and pressure solution seams (dark streaks down the left side of the photo). For Figures 2.7-2.13, stratigraphic up is towards the top of the page unless otherwise noted. FIGURE 2.8. Representative samples of the Sbl subfacies: a) outcrop of limestone lenses (Sbl subfacies) and calcareous shale (Scs subfacies) near Quarry 6 (Camp, 1980). b) Partially articulated ichthyosaurs in the Visitor's Quarry at BISP. Photo shows ribs and vertebrae (arrow, field of view about 20 feet), c) 9607-1, acetate peel, wackestone with many nuculid bivalve shells oriented convex- up and down, d) 71095-2, acetate peel (parallel to bedding), ammonoid-nuculid bivalve wackstone. The dark spots on the right side of the photo are fills of tiny gastropods. Also note spar-filled ammonoid (left) and single bivalve shells, e) 71595-36, thin section, wackestone with thin-shelled bivalve fragments, rare foraminifera (middle of photo), recrystallized, very fine-grained matrix, and iron oxide possibly replacing pyrite (square in lower right). FIGURE 2.9. Representative samples of the Scs subfacies; a) 72995-1, purplish-weathering calcareous shale. Black fresh material is visible in exposed ledge at upper right of photo, b) 72995-1, thin section. Note very fine-grained recrystallized matrix and very small bioclasts, including whole ostracods (o), foramifera? (0, and sponge spicules (s). c) 60794-10, acetate peel. Whole and disarticulated ostracods comprise most of the bioclasts in this sample. Several sets of calcite veins are also visible. FIGURE 2.10. Representative samples of the Pel subfacies: a) 72995-19, peloidal wackestone with undulating laminations (hammer is 40.5 cm long), b) 72995-23.1, acetate peel showing alternating laminations of dark- and light-colored grains, c) 60694-15, thin section showing wackestone with peloids (dark grains), bioclasts, and a few oolites (dark grains with lighter rims), d) 72995-19, wackestone with imbricated ammonoids (lower 1/3 of photo) indicating current activity. Bioclasts fme upwards. FIGURE 2.11. Representative samples of the Pbl subfacies: a) 60794-9, pelleted brachiopod wackestone, bedding plane view, b, c) 60794-9, thin section showing brachiopod shell fragments and pellets (compare with 2.10c for contrasting nature of peloids and pellets), d) 60794-7m, bioclastic wackstone. Note gastropod cross-section, bioclasts, and light-colored intraclast in lower right comer with micritized foraminifer. FIGURE 2.12. Representative samples of the O facies: a) 70795-lS, oolitic crinoid-brachiopod grainstone. b) close-up view of (a), showing faint laminations, c) 60794-13, polished slab showing coarse, poorly sorted bioclasts included brachiopod shells and crinoid columnals. d) 60794-15, thin section, showing brachiopod and echinoderm fragments (upper left), oolites (upper right), uniserial foraminifer (lower middle), and other grains. Matrix is lacking, e) 70795-15, acetate peel, brachiopod packstone immediately overlying 60794-15 (round white patches are imperfections in the peel). FIGURE 2.13. Representative samples of the F facies: a) 60994-17 through 21, gastropod-brachiopod-oyster wackestone. Asterisk marks the sole of bed 60994-21, which contains large horizontal trace fossils and oysters, b) close-up of large, articulated oysters located at the * in (a) (pencil is about 13 cm long), c) float block, large gastropod (scale in cm), d) 60994-IS, burrow-mottled outcrop (compare light and dark areas at top and bottom of outcrop), e) 60994-19, acetate peel, wackestone. Note thick brachiopod shell, echinoderm debris (round and blocky fragments), and fme-grained muddy mauix. 145

^ Major Elements and TOC 1

X S4^4^n

«g 3 •+ • •2 TA • DtocTOC E • +Mg *A1 oa 2 • APe M < I

Sla Sbl Scs Pel Pbl O

Facies

Major Elements and TOC 2

= 4 a 3E . "TOC i.A I® ^ -DNa i 3 $ ^ ^ X • xp "3 *^ , •S 2 - +K

u ^ '

Sla Sbl Scs Pel Pbl O F

Facies

FIGURE 2.14. Log of TOC and major element concentrations plotted by facies. Averages by lithofacies of elemental concentrations are shown in these graphs. See also Table 2.5. 146

Minor Elements and TOC 1 4.5

T 3.5 O ^ X ''TOC Oi e X X O ^ Xsc

IS 2.5^ • • ^ • H g DMH na O Xsr BD 1-5 ~ . +Y

i..$ o 6r-Zr i 0.5 X I$ ^ ~J, - " < * ® X X OLa -0.5 Sla Sbl Scs Pel Pbl

Facies

S Minor Elements and TOC 2 g" Otoc a O +V r O I 3 O O o ^

-3 M 2 • X i ^ Xcu XZn il l X A i 8 S < X ^ ^ °Ba

Sla Sbl Scs Pel Pbl O

Facies

FIGURE 2.15. Log of TOC and minor element concentrations plotted by facies. Averages by Iithofacies of elemental concentrations are shown in these graphs. See also Table 2.5. FIGURE 2.16. Trends in fossil abundance for taxa sampled in Section 1 of Hogler (1992a). Lithofacies are displayed in stratigraphic order with the youngest on the right. Sampling was constrained by outcrop exposure and was discontinuous, so lines connecting abundances do not necessarily reflect actual trends between samples. Taxa used in ihis figure are listed in Appendix C. See Hogler (1992a) for detailed locality and sample infonnation. Key to bivalve (Biv) trophic groups: i/dp=infaunal deposit feeder, p/su=planktic suspension feeder, e/su=epifaunal suspension feeder, and i/su=infaunal suspension feeder.

^ Gastropods • Brachiopods s\\ NNV • Biv i/dp AW ASSV vSSSW • Biv p/su vSNSWV C3Biv e/su sSNSSSWV^:: SSNNNNNW^:? E Biv i/su \NN\\\\S\vX: OvWSSNNNNvJ O.S\\\\S\Vs\\ • Nautiloids vSWSSNNWWW vSWSWSNNNSSNV Q Ammonoids Ov\\S\S\\\\\S\\>C »w\\\SSSN\\SN\\SNN vSNVSNSSWWWSSNV vSN\SNNNNNS\\\\\S\ vSS\SSSS\S\\NVS\SS\ s\\S\S\NN\\N\\\\Vv\V vS\\\SSN\\S\N\N\\\\NV vS\SS\S\\N\\SSNSN\SSSV K\\\N\N\\\\\\\\\\\\\VS\

vNVsNNVSNV

Sla Sbl Scs Sbl-Pel Pbl-O

Lithofacies -ti. -J Mostly whole fossils Mostly bioclasts Partly articulated bones Fecal pellets HHMHBHB Clotted fabric Burrows/traces Peloids - Ooids —— Primary sed. structures ^ —— Fine-grained sediment

I SWB 'FWB Scs Sbl Sla Pel Pbl O F

Sea level Ooid Inner shelf shoals

FWB Pbl

SWB Pel

Sla Sbl FIGURE 2.17. Lithofacies plotted by water depth (lower diagram) Scs and trends in sedimentary and taphonomic observations (top diagram). The width of the line in the top diagram indicates the relative expression of the feature by facies. FIGURE 2.18. Ammonoid morphotype abundances by lithofacies. Abundance data from section 1 of Hogler (1992a). Morphotype ecology information from Westemiann (1996). Taxa listed in legend; none were present in the Pbl, O, or F lithofacies. Line drawings of lateral and ventral views adapted from photographs on plates 1, 2, and 5 of Silberling (1959); figured taxa are indicated in bold except for the planktic drifter category, where the taxon is Tropiceltitesl densicostatus Silberling 1959.

•nekton 160 Klamathites macrolobatus Klamathites sp

140 •planktic drifter

120 Tropicehites columbianus ?juvenile ammonite

100 •demersal/planktic drifter/nekton

80 cf. Mojsisovicsites sp. Ik Mojsisovicsites sp. I «0 67 ^ 60 •vertical planktic migrant

Arcestes sp. 40 cf. Guembeliles sp. m Guembeliles jandianus Guembeliles sp. 20 Juvavites sp. Tropites crassicostatus n M Tropites laliumbilicalus Sla Sbl Scs Sbl-Pel Tropites subquadratus Lithofacies Tropites sp. 150

Scenario 1

Normal-slow Bone lags sedimentation OMZ stagnant basin

Shelf Most intense oxygen depletion BISP environments Basin

Upwelling-fovorable wind (soutn tor western Nonn America) Scenario 2 Production surface water moves locus L*J offshore —

Rapid sedimentation of biogenic particles

OMZ intensified by upwelling

Shelf

Basin BISP environments

Upwelling-favorable wind (south for western North America) Scenario 3 Production surface water moves locus offshore

Seasonal influx of terrigenous sediment Rapid sedimentation helps preserve seasonal of biogenic particles • upwelling signal (if ooid shod was not yet S=3 developed) OMZ intensified by upwelling

Shelf Basin BISP environments

FIGURE 2.19. Scenarios for deposition of the Luning Formation based on evidence from BISP. Scenario 3 only applies if the ooid shoal was not fully developed (possibly for the lower half of Luning deposition at BISP). See text for details. 151

TABLE 2.1. Preservable macrofaunal characteristics of eutrophic ecosystems, adapted from Brasier (1995), Hallock and Schlager (1986), and Parrish (1995). Oligotrophic (normal marine) ecosystems do not usually show these characteristics. The three rightmost columns show macrofaunal characteristics for sediments that have been interpreted as ancient upwelling deposits. Other sources used in the body of the table include Allmon (1993), Ailmon et al. (1996), Brongersma-Sanders (1948), Brongersma-Sanders (1957), Hallock and Schlager (1986), Hiller (1994), Lescinsky (1996), Parrish and Gautier (1993), and Seibold(1982). + = present; - = absent: ? = unknown

Macrofaunal characteristic Eutrophic Permain Sharon Pliocene ecosystems Phosphoria Springs sediments, Fm. Member* SW Florida Abundant organisms + (r-selected taxa, large n) High dominance + +? Low diversity + (benthos) Mass mortality (unstable + populations) Non-symbiotic suspension + feeders common Marine vertebrates common + + + Lack of symbionts (esp. + + + zooxanthellate corals) Cool-water taxa in low- latitudes (unless upwelling) Depauperate benthos in + oxygen-minimum zone (nearshore fades deposits) Organic-rich fecal pellets + + + Bioerosion common + 9 9

'Cretaceous Pierre Shale TABLE 2.2. Estimates of sedimentation rate for ammonoid Zones in the Luning Formation at West Union Canyon. Ammonoid Zones were assumed to have equal duration within each stage; Zone durations were interpolated from Figure 9 of Gradstein et al. (1994); age of the Camian-Norian boundary (220.7 ± 4.4 Ma) is also from Gradstein et al. (1994). Thickness A interpolated from Plate 11 of Silberling (1959). Thickness B from Fossil Hill (Welleri Zone) and Boundary Hill (Macrolobatus and Kerri Zones) columnar sections of this dissertation (see Appendix A).

British West Union Lower Upper Duration Thick­ Thick­ Rate A Rate B Lithofacies Columbia Canyon Local bound bound my ness A, ness B, cm/ky cm/ky Regional Ammonoid Ma Ma m m Ammonoid Zone Zone

Kerri Guembelites 220.7 ± 219.2 1.5 30.5 22 2.0 1.5 Scs-Pel 4.4 Macrolobatus Macrolobatus 222.2 220.7 ± 1.5 41.1 30 2.7 2.0 Sbl-Scs 4.4 Welleri Schucherti 223.7 222.2 1.5 38.1 38 2.5 2.5 Sla TABLE 2.3. Summary of lithofacies, Luning Formation, West Union Canyon. In the "Color" column, W = weathered color and F = fresh color.

Fades Subfacies Lithologies Bed Color Sedimentary Characteristic Modes of Interpretation thickness structures fossils preservation s. Sla, shaly wacke- thick bedded tan, brown, gray. none visible infaunal, sus­ internal and Distal storm Shaly Silty, brown stone and green (W); dark pension-feeding external molds deposits, just fossili- limestone & packstone. gray-brown (F) bivalves, gastro­ below storm ferous argillite argillite pods wave base limes­ Sbl, shaly micrite. thin - thick tan or blue-gray burrows?. ammonoids. recystallization, Offshore, below tone Silty, black wackestone. bedded (W): black (F) bivalve gastropods, nuculid external molds storm wave base. limestone packstone laminations and halobiid dysoxic benthic bivalves. conditions ichthyosaur bones Scs calcareous thick - very tan, purplish bivalve ostracods, halobiid recrystallization. Offshore, below Calcareous shale thick bedded (W); black (F) laminations bivalves, external molds storm wave base. shale radiolarians, sponge dysoxic benthic spicules conditions P. Pel, peloidal medium blue-gray. flat and bioclasts (peloids). recrystallization Medial storm Pel- Peloidal wackestone bedded, with brown, gray. swaley ammonoids. deposits. bio- bioclastic and pack­ some sharp green (W); dark- lamination. halobiid bivalves. between stonn clastic limestone stone basal gray, black (F) normally ostracods. and fair-weather lime­ contacts graded beds, echinoderm wave base stone imbrication fragments Pbl, pelletal thick bedded light blue, gray none visible fecal pellets, recrystallization Energetic Pelleted wackestone (W); gray, black bioclasts. (fecal pellets not environment. bioclastic and pack­ (F) brachiopod shells compacted) early cemen­ limestone stone tation O, Oolitic bioclastic oolitic thick bedded gray, dark gray small- crinoid and recrystallization. Carbonate limestone (no packstone (W); dark-gray. medium scale brachiopod frag­ micrite envelopes shoals, ooids subfacies) and grain- black (F) crossbeds. ments indicate ener­ stone normally getic conditions graded F, Fossiliferous wackestone medium - dark gray (W); burrows. brachiopods. reciystallization Back-shoal, limestone (no and pack­ thick bedded dark gray, black bioclast gastropods. quiet-water subfacies) stone (F) laminations echinoderms. environment ostreids 154

TABLE 2.4. Orientations of 441 Halobia sp. shells. Shells were counted on both the tops and bottoms of slabs, as indicated by the "Side" field. Only shells with discernible orientations were counted.

Sample # Piece Side Umbo Umbo Articu­ Notes convex convex lated down up Oriented slabs 72295-24 A top 0 2 many shells uncounted; shale 72295-24 B top 0 1 many shells uncounted; shale 72795-27b B bottom 2 3 72995-1.5a A bottom 1 0 72995-1.5a B bottom 0 3 72995-1.5a C 7 3 3 72995-23.5a A top 6 5 72995-23.5a A bottom 3 1 72995-23.5a B top 3 4 72995-23.5a B bottom 11 5 72995-23.5a D top 10 7 72995-23.5a D bottom 15 21 72995-24 A bottom I 3 72995-24 C bottom 0 1 72995-25b top 7 6 72995-25b bottom 3 5 72995-27b top 5 2 72995-27b bottom 1 0 960718-23b23.5 A top 6 11 960718-23b23.5 A bottom 8 9 960718-23b23.5 B top 4 4 960718-23b23.5 B bottom 7 5 Total for oriented slabs 96 101 155

TABLE 2.4. (continued)

Sample # Piece Side Umbo Umbo Articu- Notes convex convex lated down up Unoriented slabs 23b I 5 8 23b 2 7 6 606941-24 1 of2 A 1 3 7 606941-24 1 of 2 A 2 1 2 606941-24 1 of 2 B 1 3 1 606941-24 2 of 2 A 1 10 10 606941-24 2 of 2 A 2 3 1 1 ?matched pair, not articulated 606941-24 2 of 2 B 1 12 6 72095-2 1 3 2 72095-2 2 1 0 72905-2f I 1 0 72905-2f 2 I 0 72995-1 Of A 1 10 7 72995-1 Of A 2 9 6 72995-1Of B I 1 1 72995-1 Of C I 8 5 72995-1 Of C 2 2 0 72995-1 Of D 1 0 3 72995-IOf D 2 1 1 72995-23.5b 1 6 12 72995-23.5b 2 5 3 1 possible articulated specimen 72995-24 B 1 1 1 72995-24 D I 0 3 72995-24 E 1 0 3 72995-28.5 A 1 1 3 shale 72995-28.5 A 2 1 6 shale 72995-28.5 B 1 2 1 shale 72995-28.5 C 1 0 1? shale 72995-28.5 D 1 0 3 shale Campground C A 1 1 2 Campground C A 2 3 8 Campground C B 1 6 4 1 ?matched pair, not articulated Campground C B 2 1 0 Campground C C I I 2 Campground C C 2 1 1 72995-23.5a C 8 8 I matched shells close by but not articulated Total for unoriented slabs 118 126 4 possibly matched shells

Total Shells Counted 441 TABLE 2.S. Geochemical results grouped by lithofacies, excluding 12 elements that had concentrations below analytical detection limits. See Appendix B for complete results of individual samples. V, Cr, Zr. and Ba concentrations are minimum estimates.

Litho­ # of Samples Detection TOC Na Mg Al P K Sc V Cr Mn Fe facies limit 0.01 0.01 0.01 0.01 0.01 0.5 2 1 2 0.01 roc ici' Units \\1 % wt % \\1 % \\1 % wt % \vt% ppm ppm ppm ppm \M %

Sla 2 4 Mean 0.14 0.12 0.36 1.23 0.07 0.21 3.9 21 18 1325 1.51 Max 0.19 0.17 0.47 1.67 0.09 0.34 5.5 25 23 1590 1.91 Min 0.09 0.06 0.22 0.74 0.05 0.05 2.4 13 14 871 1.18

Sbl 5 6 Mean 0.27 0.10 0.69 1.18 0.04 0.16 2.4 46 16 417 0.97 Max 0.38 0.23 l.ll 2.33 0.05 0.47 5.4 181 18 796 1.50 Min 0.18 0.05 0.30 0.43 0.03 0.02 0.8 9 10 204 0.47

Scs 4 8 Mean 0.66 0.23 0.61 2.27 0.04 0.46 4.6 73 27 292 1.40 Max 0.85 0.54 1.08 5.31 0.12 1.02 10.4 136 45 563 3.30 Min 0.46 0.12 0.35 0.85 0.03 0.22 2.2 21 13 157 0.68

Pel 3 3 Mean 0.14 0.06 0.36 0.39 0.03 0.09 0.9 20 8 145 0.32 Max 0.24 0.09 0.39 0.67 0.03 0.15 1.5 32 10 174 0.47 Min 0.08 0.03 0.34 0.20 0.02 0.03 0.5 8 6 105 0.19

Pbl 1 I n/a 0.01 0.16 0.26 0.57 0.04 0.24 1.6 12 8 230 0.13 n/a n/a

O 1 1 n/a 0.06 0.04 1.03 0.44 0.04 0.20 1.7 13 11 357 0.96 n/a n/a

F 3 3 Mean 0.13 0.04 0.38 0.55 0.02 0.24 1.4 12 11 188 0.42 Max 0.24 0.04 0.62 0.74 0.03 0.33 1.5 16 16 252 0.72 Min 0.04 0.03 0.26 0.34 0.01 0.15 1.2 8 7 129 0.13 TABLE 2.5. (continued) Lilho- Hof Detection Ni Cu Zn Sr Y Zr Ba La fades Samples limit 1 0.5 0.5 0.5 0.1 0.5 1 0.5 TOC ICP Units ppm ppm ppm ppm ppm ppm ppm ppm

Sla 2 4 Mean 23 25.5 39.2 388.3 10.9 12.8 90 7.1 Max S8 79.9 56.6 601.0 11.7 19.8 117 10.0 Min 8 4.8 27.1 144.0 10.3 9.7 51 4.6

Sbl 5 6 Mean 15 9.6 66.0 1175.8 4.9 18.4 50 4.2 Max 19 16.9 128.0 1480.0 10.2 36.2 120 7.5 Min II 4.9 42.1 633.0 2.6 7.1 II 2.6

Scs 4 8 Mean 34 17.1 111.9 1424.9 10.0 27.2 123 7.1 Max 89 32.7 242,7 1783.3 13.7 52.8 257 15.2 Min 14 8.4 37.6 649.0 4.6 16.8 62 3.3

Pel 3 3 Mean 10 3.4 36.6 2876.7 2.9 11.0 36 1.9 Max 11 4.5 46.2 3270.0 4.9 12.3 58 2.5 Min 8 2.8 26.0 2120.0 1.6 9.2 17 1.5

Pbl 1 1 n/a 8 4.5 14.4 736.0 14.6 16.4 12 5.9 n/a n/a

O 1 1 n/a 14 4.8 9.8 368.0 8.3 7.1 14 5.0 n/a n/a

F 3 3 Mean 27 8.3 20.7 677.0 3.3 8.0 10 2.6 Max S8 15.9 23.5 793.0 3.8 8.5 14 2.9 Min 9 3.2 18.9 463.0 2.9 7.4 8 2.3 TABLE 2.6. Spearman rank correlation coefficients for log of TOC and of major and minor elements shown in Table 2.5. Coefficients were calculated using Systat (listwise deletion) on 18 samples that were subjected to both TOC and ICP analysis (Tables B.I and B.2). Pairs with coeiTlcients > 0.830 are TOC/Zn, Al/Sc, Sc/La, Cr/Fe, Cr/Cu, and Ni/Cu.

Log TOC NA MG AL P K SC V CR MN FE NI CU ZN SR Y ZR LA

TOC 1.000

NA 0.680 1.000

MG 0.251 -0.054 1.000

AL 0.768 0.801 0.041 1.000

P 0.527 0.687 -0.055 0.585 1.000 K 0.501 0.659 -0.036 0.752 0.429 1.000 SC 0.588 0.809 -0.017 0.929 0.653 0.753 1.000 V 0.711 0.648 0.117 0.779 0.396 0.656 0.745 1.000

CR 0.698 0.661 -0.053 0.757 0.581 0.553 0.742 0.591 1.000

MN -0.023 0.206 -0.241 0.284 0.401 0.020 0.419 -0.050 0.484 1.000

FE 0.484 0.476 -0.097 0.567 0.540 0.230 0.611 0.333 0.870 0.762 1.000 N1 0.620 0,295 0.171 0.564 0.173 0.577 0.434 0.526 0.721 0.027 0.409 1.000

CU 0.697 0.547 0.014 0.811 0.478 0.695 0.700 0.579 0.874 0.293 0.617 0.875 1.000

ZN 0.859 0.621 0.237 0.687 0.430 0.322 0.548 0.760 0.653 -0.001 0.451 0.504 0.600 1.000 SR 0.321 0.033 0.218 -0.161 -0.219 -0.137 -0.343 0.147 -0.200 -0.808 -0.444 0.049 -0.153 0.366 1.000

Y 0.297 0.777 -0.161 0.642 0.531 0.691 0.778 0.544 0.503 0.293 0.368 0.199 0.388 0.278 -0.347 1.000

ZR 0.617 0.830 0.081 0.769 0.390 0.577 0.724 0.796 0.533 -0.013 0.279 0.390 0.527 0.702 0.166 0.612 1.000

BA 0.659 0.819 0.103 0.806 0.552 0.578 0.831 0.776 0.644 0.172 0.528 0.290 0.521 0.686 0.059 0.544 0.803

LA 0.311 0.654 -0.222 0.765 0.664 0.639 0.875 0.480 0.621 0.542 0.591 0.261 0.588 0.239 -0.610 0.770 0.538 TABLE 2.7. Geochemical results for the tuning Fm. at West Union Canyon compared with a) geological reference materials (nos. 8-12), b) modem marine plankton and diatom ooze (nos. 13,20), c) samples from ancient upwelling deposits (nos. 14-17), eind d) samples from deposits with high productivity enhanced by terrestrial mnoff (nos. 18-19). TOG in weight percent; all other concentrations in ppm.

Description TOC Na Mg A1 P K Sc V Cr Mn Fe Ni Cu Zn Sr Y 'L\ Mo Cd Ba La 1. Sla facies 0.14 1163 3563 12263 713 2075 4 21 18 1325 15100 23 26 39 388 11 13 2 <1 90 7 2. Sbl facics 0.27 1033 6917 11750 450 1633 2 46 16 417 9750 15 10 66 1176 5 18 7 <1-10 50 4 3, Scs facies 0.66 2319 6088 22671 706 4590 5 73 27 292 14021 34 17 112 1425 10 27 4 <1-9 123 7 4. Pel facies 0.14 567 3633 3867 267 867 1 20 8 145 3233 10 3 37 2877 3 11 2 <1-2 36 2 5. Pbl facies 0.01 1600 2600 5700 400 2400 2 12 8 230 1300 8 5 14 736 15 16 1 <1 12 6 6. O facies 0.06 400 10300 4400 400 2000 2 13 11 357 9600 14 5 10 368 8 7 3 <1 14 5 7. F facies 0.13 350 3833 5483 200 2433 1 12 11 188 4217 27 8 21 677 3 8 3 <1 10 3 8. Limestone 964 4463 12648 528 3403 3 24 15 682 20 10 22 545 14 35 2.1 50 13 9. Crinoldic limestone 371 16344 1693 218 664 27 5 77 11 29 267 7 35 26 5 10. Argillaceous limestone 2893 2171 5927 349 2075 29 16 1549 11 6 41 1180 16 86 II. Carbonaceous shale 6900 48851 71442 218 11538 139 123 465 27277 122 53 76 54 17 146 5 0.2 820 35 12. Average shale 0.20 88000 700 130 90 850 68 45 95 2.6 0.8 580 13 Marine plankton 7600 3 2 8 11 no 2 12 80 14. Calcareous mudstone' 2.41 5490 4885 47205 28890 15274 10 227 1280 124 19303 115 68 370 237 249 8 18 277 148 15. Argillaceous limestone* 0.89 519 60189 22914 698 9629 4 192 135 132 8183 80 14 590 104 10 9 57 90 10 16. Calcareous mudstone'' 2.50 8309 5971 18204 1004 6392 4 126 65 155 9722 56 22 70 451 8 79 10 7 168 9 17. Phosphatic shale' 16.00 13503 7780 35721 16889 9795 6 370 315 155 16296 240 91 267 605 23 39 26 8 246 17 18. 'Posidonia Shale' 2374 12665 49745 873 14112 160 66 697 32382 91 61 206 1355 23 2 255 19. Peterborough Member'' 5.11 3784 8564 90705 991 22413 20 140 166 194 33571 56 32 110 279 32 167 6.6 329 46 20. Namibian shelf diatom 9.35 15400 108 68 68 53 ooze 'Meade Peak Member, Phosphoria Formation (Periman) *"8150 shale and porcelanite, l.o\ver calcareous-siliceous member, Monterey Formation (Miocene) 'Carbonaceous marl member, Monterey Formation (Miocene) ''Oxford Clay (Jurassic) Rerercnces; Sample number (see references for details); 1-7 This dissertation 1-4, 7,12-13.18-20 Composite 15 P-79 8-11 Potts et al., 1992 5 Field number 60794-6 16 KG-11 12-13 Brumsack, 1986, Wedepohl, 1969-78 6 Field number 70795-15 17 KG-2 14-15 Medrano and Piper, 1992 8 Kll 16-17 Piper and Isaacs, 1995 9 CCH-I 18 Brumsack, 1991 10 SRM lb 19 Norry et al., 1994 11 SARM41 20 Calvert and Price, 1983 14 P-165 160

TABLE 2.8. Expected characteiistics of the scenarios for Luning deposition and observed features. See text for details. + = characteristic expected, - = not expected, +/- = may or may not be expected, ? = unknown Scenario 1 Scenario 2 Scenario 3 Observed Enhanced Marine Seasonal features in the OMZ upwelling runoff and Luning Fm. at Feature upwelling BISP Depauperate benthic fauna in + + + + OMZ fades Organic-rich sediments in OMZ + + + (+) facics

The most oxygen-depleted facies + - - - are in the center of the basin

Ichthyosaur bone in lags or + - - - concentration deposits

The most oxygen-depleted facies - + + + are offset from the center of the basin Enriched in elements associated -/+ + + (+) with marine plankton (e.g., P, Zn, Cu, Ni) +/- Cool-water taxa at low latitudes - + +/-

Lack of hermatypic corals and - + + + calcareous algae + + Abundant suspension feeders - + + + Abundant terrigenous debris +/- - (when clastic transport occurred. (lower half not during oolite formation) of section) +/- + 9 Terrestrial component of organic - matter TABLE 2.9. Distribution of marine vertebrates reported in the literature from Triassic sediments of the Mesozoic marine province. Marine vertebrates are found in the region throughout the Triassic. System Locality Formation Marine reptile Abundance & References Series order Completeness Stage Triassic Pilot Mountains, Luning Ichthyosauria ?, many bones Stanley, 1979 Upper Nevada Formation found in float Norian Camian Humboldt Range, Natchez Pass Thalattosauria 1 specimen Storrs, 1991 Nevada Formation Shoshone Mountains, Luning Ichthyosauria 37+ well-preser­ Camp, 1980; Hogler, 1992b; Nevada Formation ved specimens Silberling, 1959 and fragments Middle New Pass Range, Favret Ichthyosauria 3 well-preserved Callaway and Massare, 1989; Anisian Nevada Formation specimens Merriam, 1908 Tobin Range, Nevada Favret Ichthyosauria 1 fragment Sander etal., 1994 Formation Augusta Mountains, Favret Ichthyosauria, 8+ well- Merriam, 1910; Sander and Nevada Formation Nothosauria, preserved Bucher, 1990; Sander et al., Plesiosauria*, specimens and 1997; Sander et al., 1994 Thalattosauria ftngments Humboldt Range, Prida Formation Ichthyosauria, 7+ well- Mazin, 1986; Merriam, 1908; Nevada Sauropterygia^ preserved Motani, 1997; Silberling and specimens and Nichols, 1982 fragments Lower southeast Idaho Thaynes Ichthyosauria 1 fragmented Massare and Callaway, 1994 Formation specimen Humboldt Range, Prida Formation Sauropterygia^ 1 fragmented Mazin and Bucher, 1987; Spathian Nevada specimen Motani, 1997 *Pistosauridae ^Omphahsaurus CHAPTER 3

PALEO-UPWELLING AND THE DISTRIBUTION OF MESOZOIC

MARINE REPTILES 163

INTRODUCTION

In this chapter, I examine the distribution of extinct marine reptiles relative to upwelling deposits and predicted upwelling zones. Marine reptiles have been proposed as ecological equivalents of modem whales (Parrish and Parrish 1983), which feed heavily in upwelling areas. Additionally, whale evolution has been linked to tectonic and climatic events that might have increased upwelling (Fordyce 1980; 1991). Marine reptiles might have had similar ties to upwelling areas to provide food. If this were the case, then marine reptile fossils would be distributed non-randomly relative to predicted upwelling.

Modem Whale Distribution and Feeding

Today, whales, dolphins, and porpoises inhabit a variety of food-rich habitats, nearly all associated with upwelling, including shallow seas, thermal fronts, and open- ocean divergences (Norris 1983). High productivity in shallow seas can result from upwelling or from nutrient recycling. Whales feed in all major coastal upwelling areas

(e.g., Townsend 1935; Gallardo et al. 1983; Reilly and Thayer 1990; Tershy 1992), as well as on shelves that are productive through nutrient recycling, such as the Bering Sea and the westem North Atlantic Ocean (e.g., Baretta and Hunt 1994; Woodley and Gaskin

1996). Thermal fronts are characteristic of geostrophic currents such as the Gulf Stream, which has highly productive eddies and meanders that enhance shelf-edge production

(Olson et al. 1994; upwelling associated with westem boundary currents). Sperm whales have been observed to congregate along the shelf edge in the Gulf Stream, probably in relation to the elevated productivity there (Waring et al. 1993). Many larger whales. 164 such as blue whales and sperm whales, feed and are sighted in open-ocean upwelling areas (Reilly and Thayer 1990; Jaquet and Whitehead 1996).

Cetacean dependence on high-productivity areas is further suggested by whale abundance trends over a 3-year period in the Gulf of California (Tershy et al. 1991).

During the 1983-84 El Nifio-Southem Oscillation event, primary production decreased dramatically in the Eastern tropical Pacific upwelling areas but was unaffected in the

Gulf of California, where cetacean sightings increased. Cetacean sightings decreased after the ENSO, suggesting that perhaps the Gulf of California acted as a refuge for food for mobile pelagic animals (Tershy et al. 1991).

Some baleen whales are observed to feed only when prey reach certain densities

(Dolphin 1987; Forcada et al. 1996). The stispension-feeding habits of baleen whales necessitate prey populations that are dense enough to make foraging feasible.

Consequently, baleen whales are usually found in areas where prey are dense—typically areas of upwelling and fronts (Forcada et al. 1996). Similarly, odontocete whales are associated with high productivity. For example, in 1992-93, sperm whale density in the southeastern Pacific was most strongly associated with the distribution of their prey, namely cephaiopods. In this case, the sperm whale distribution was most closely associated with a measure of secondary rather than primary productivity (Jaquet and

Whitehead 1996). The sperm whales were not associated with measures of secondary productivity, such as zooplankton, at small spatial scales (less than 80 nautical miles), but this result probably reflects the spatial and temporal lags between zooplankton and cephalopod populations (Jaquet and Whitehead 1996). Killer whales follow the 165 distribution of their prey (smaller marine tetrapods and fish) (Simila et al. 1996), and would thus be associated with production at high trophic levels. Killer whales are voracious predators and can eat numerous prey (e.g., the stomach of a 7.3-meter-long killer whale contained remains of 13 porpoises and 14 seals; Slijper 1979).

In summary, observations of whale feeding and distribution ( Townsend 1935; charts in FAO Department of Fisheries 1972) show that whales are strongly associated with areas of high productivity for a significant portion of their lives.

Marine Reptile Ecology

Marine reptiles likely had similar ecologies to modem whales (Petkewich 1982;

Parrish and Parrish 1983), particularly toothed whales (Massare 1987). All marine reptile taxa show morphological similarity to cetaceans (and other marine tetrapods) that relates to streamlining of the body shape, including reduction of limbs, reduction of vertebral articulations, and enlargement of the lumbar region (Gaskin 1982). Beyond these streamlining changes, marine reptiles show varying degrees of morphological similarity to whales. Ichthyosaurs are perhaps the most similar to whales, with their short necks, paddle-like limbs, lunate tail, and elongated rostrum. Plesiosaurs retained long necks

(except for pliosaurs), long tails, and long paddles, while mosasaurs retained long tails.

Modem whales use highly specialized features to utilize dense food resources, namely baleen plates in baleen whales and echolocation in toothed whales. Marine reptiles do not have the mammalian features of the skull that would allow suspension feeding (Collin and Janis 1997) or the generation and reception of a sonar beam (Taylor

1987). Morphological adaptations that are related to feeding are more difficult to 166 determine for marine reptiles, but several species of ichthyosaurs had very large eyes that could have aided food capttire (Collin and Janis 1997).

Stomach contents found in association with marine reptile bones suggest feeding habits similar to many of today's toothed whales (see table 1 in Massare 1987). Fish vertebrae, cephalopod booklets, and remains of larger vertebrates, including pterodactyls, smaller ichthyosaurs, and flightless birds, are the most common constituents among the few reported occurrences of stomach contents (Massare 1987). Marine reptile teeth have similar morphologies and thus inferred functions as odontocete teeth. Based on tooth morphology, marine reptiles can be grouped into several feeding guilds that define prey groups such as sofl-bodied prey, fish, large vertebrates, and shelled prey (Massare 1987).

Baleen whales are the largest icnown tetrapods, attaining lengths of up to 30 meters, while odontocete whales attain lengths of up to 15 meters (Lalli and Parsons

1993). The largest known marine reptiles include the pliosaurs Liopleurodon ferox

Middle Jurassic) and Kronosaurus queenslandicus (Lower Cretaceous), the ichthyosaur

Shonisaurus popularis (Upper Triassic), and the mosasaur Mosasaurus hqffmami (Upper

Cretaceous), all of which attained lengths of about 15 meters (Quinet 1970; Kosch 1990;

Martin 1991; Long 1998). Large body size increases an animals' food requirements but also allows the animal to retain heat more efficiently by lowering the surface area to volume ratio (Ricklefs 1990). Histological studies of ichthyosaur bone from two specimens showed remodeled bone, which indicates resorption, and woven-fiber tissue, which can indicate a high growth rate that suggests endo- or giganto-thermy (de Buffrenil 167 and Mazin 1990). It is possible that some marine reptiles had high food requirements related to metabolism and a large body size.

Upwelling Characteristics

As described in Chapter 1, modem upwelling areas are typically sites of high biologic productivity with unique sedimentological, biological, and geochemical characteristics. These features can be preserved in the geologic record and used to identify sites of ancient upwelling. For example, marine organic-rich, oil-prone sediments are almost exclusively found in predicted upwelling areas (Parrish 1995).

Other sediments commonly found in upwelling areas include phosphorite, biogenic silica, and glauconite, especially all three in association (Parrish 1995, see also Chapter 1).

Abundant remains of large, marine predators are considered characteristic of upwelling zones (Parrish and Gautier 1993; see Chapter 1 for further discussion of upwelling biotas;

Brasier 1995).

Overview of the Studv

This study uses paleogeographic analysis to assess the relationship between marine reptile fossil distribution and predicted upwelling areas for the Mesozoic.

Whereas several authors have suggested that food availability could have influenced marine reptile distribution (McGowan 1972; Nesov et al. 1988; Hogler 1992b; Gasparini et al. 1996; Collin and Janis 1997), only one study has investigated the relationship.

Parrish and Parrish (1983) compared marine reptile localities and predicted upwelling and found that for all Mesozoic stages together, 67 % of reptile fossils were located in or near predicted upwelling areas, which occupied about 25 % of the shelves. The present 168 study expands on the Parrish and Parrish (1983) study by including more reptile taxa

(Parrish and Parrish, 1983, focussed on sauropterygians, while this study includes ichthyosaurs, mosasaurs, and crown-group sauropterygians), updated palgeographic reconstructions for ten stages of the Mesozoic, and data for upwelling-related lithologies including phosphorite, biogenic siliceous sediment, and glauconite. 169

METHODS

Database Design and Assembly

In order to address Mesozoic marine reptile distribution, I assembled a relational database of fossil reptile localities. For the database, I chose 3 large clades of fully aquatic reptiles: ichthyosaurs, mosasaurs, and plesiosaurs. The vast majority of taxa from these orders are fully marine, but 3 pliosaur genera have been found in lacustrine and fluvial sediments and are interpreted as freshwater dwellers {Bishanopliosaurus,

Leptocleidus, and Sinopliosaurus) {Gds^^ni 1992; Cruickshank 1997). Data were collected for these taxa but were not included in the analyses discussed here. Nothosaurs, placodonts, other ancestral sauropterygians, and thalattosaurs were excluded because these taxa are thought to have been either littoral dwellers or capable of moving on land

(Tarlo 1967; Benton 1979; Rieppel 1989; Nicholls and Brinkman 1993). Marine crocodiles and turtles were also excluded because they were not as abundant as the three groups that I focussed on in this study.

The database was designed as a series of 4 tables related by key fields (Figure

D.l, Table D.l). The key fields, or keys, establish the relations among table records and uniquely identify each record. The relational design reduces data redundancy, thereby increasing accuracy, efficiency, and querying ability (see Hernandez 1997, for more information on relational database design). The relational design also allows one to keep track of localities and the individual taxa present at each locality (taxon-localities). To facilitate data entry, a few fields, such as the reference field, were not designed relationally. Because of the variety of data collected and the varying data quality, some 170 fields for some records have null values. Data were initially entered into Microsoft Excel and then exported into Microsoft Access and Maplnfo Professional®.

Each record in the database consists of a single marine reptile fossil locality. The basic information required to create a record includes locality name, age (at least to series), and taxon. Where possible, stratigraphic, taphonomic, and associated faunal data were entered for each record. Each locality name was assigned present-day latitude- longitude coordinates using either reported coordinates, estimations of coordinates from locality maps, or estimations of coordinates using other maps and the GeoNet names server (http://164.214.2.59/gns/html/index.html) or the USGS Geographic Names

Information System (http://www-nmd.usgs.gov/www/gnis/). Data from broad geographic regions, such as a county or small province, were included but given higher error ratings (Table D.2). Geographic coordinates were converted to decimal degrees with a precision of 1/10 degree (Table D.3). Degrees were rounded following the method of Sokal and Rohlf (1995, p. 15). Each locality was assigned to a time interval for plotting on one of the ten paleogeographic reconstructions (Tables 3.1 and 3.2).

Taxonomic schemes were constructed for each of the 3 clades, using Bardet

(1995) and recent revisions and discoveries reported in the literature (Tables E.1-E.3).

Whereas there is considerable debate about a few of the taxonomic assignments (e.g., placement of Polycotylidae, a family of Cretaceous short-necked taxa within the

Plesiosauria), the schemes provide a useftil starting place for analysis. 171

Upwelling-Associated Litholoeies

Characteristic sediment types and facies patterns are found in areas of Recent,

persistent upwelling (see Chapter 1). The sediments commonly associated with

upwelling include varying combinations of organic-rich shales, phosphorites, bedded

chert, and glauconite (Baturin 1983; Parrish 1983; Summerhayes et al. 1995). Not all

sediment types are present in every upwelling zone, and the sediments of modem

upwelling zones vary considerably. However, a facies association of biosiliceous

sediment (especially diatomaceous sediment for Cretaceous and younger deposits),

phosphorite, and organic-rich sediment strongly suggests coastal upwelling (sometimes

termed the Si-C-P association, Cook 1976). The distribution of these sediments provides

another way to assess the relationship between fossil reptile localities and ancient

upwelling zones. Therefore, in addition to the predicted upwelling areas, I plotted the

distribution of organic-rich shale, phosphorite, bedded chert, and glauconite to compare

with fossil reptile localities. Organic-rich rock data were obtained from J.T. Parrish

(unpublished data, see Parrish 1995). Localities contained organic-rich rocks that either

were interpreted as oil-prone or have hydrogen indices > 350; localities were dated to the stage level. Locality data for phosphorite, bedded chert, and glauconite for 10 stages of the Mesozoic were obtained from D. B. Rowley and A.M. Ziegler (unpublished data,

Paleogeographic Atlas project). The stages that contain these data are the same as the stages used for paleogeographic maps (see below), with the exception that Volgian data were plotted on the Kirrmieridgian map, and Induan data were plotted on the Anisian 172

map. All lithologic data were coded according to the time interval scheme shown in

Table 3.2.

Paleogeographic Reconstructions and Upwelling Predictions

Paleogeographic base maps from Scotese and Golonka (1992) were used for the

data analysis (Figures F.1-F.5; Table 3.1). Because maps are available for only 10 stages

of the Mesozoic, data were coded into three categories of increasingly finer age

resolution—series, unrestricted stage, and restricted stage (Tables 3.2 and D.2).

Unrestricted stage includes all data points reported to the stage level, regardless of

whether or not the data fall into one of the 10 mapped stages. Restricted stage includes

only those data points reported to one of the 10 mapped stages.

I digitized predicted upwelling areas onto the maps following Parrish (1982;

1995) and Parrish and Curtis (1982), modifying the predictions where necessary to reflect

revised paleogeographic interpretations. Parrish (1982) described in detail the method for

predicting ancient upwelling by modeling atmospheric circulation for past geographies.

The method assumes that in the past, like today, general atmospheric circulation was basically zonal (follows lines of latitude). The positions of the continents are then used to predict large-scale cellular features, such as monsoonal circulation, that disrupt the zonal pattern (see Parrish 1982, for discussion and examples). Upwelling areas were predicted only for areas over continental shelf because nearly all of the deep-ocean sedimentary record, particularly for the early Mesozoic, has been destroyed through subduction.

Upwelling areas were plotted where currents would diverge from a shoreline or under a 173 low pressure cell. Equatorial upwelling was plotted to include the shelf from 2° north to

2° south of the equator.

Areas in square kilometers of continental shelf and predicted upwelling zones were calculated for each map (Table 3.3). Shelf area was calculated by first generating a

2° X 2° grid of paleogeography for each map interval using the program Paleoclimate (a parametric climate model which also uses the Scotese and Golonka, [1992] reconstructions). The program assigns each grid cell to one of four paleogeographic categories: ocean, shelf, land, or highlands. The assigimient is made based on the kind of paleogeography that makes up the largest area within the cell. Thus, cells with a majority of shelf will be counted as all shelf, making up for the cells that contain some shelf but too little to be assigned to the shelf category. The grid was then divided into 2° latitude bands and the area of each band relative to the equator was calculated using the following equation:

^ sinA„-sin/l, 2(sm 1)

where Sui is the surface area between two lines of latitude u and / on a sphere of radius 1, with u representing the higher latitude and I representing the lower latitude. Next, the number of grid cells per 2° band coded by the model as shelf was multiplied by the relative area for that band. Finally, the relative area of shelf for each 2° latitude band was multiplied by the earth's circumference (40076 km) and the products summed to obtain 174 the "actual" shelf area (i.e., the shelf area at a 2° resolution. Table 3.3). Upwelling zone areas were calculated the same way, except that the upwelling zone polygons were imported into the climate model program, where they were coded as shelf.

Data Plotting Procedures

The point data (raw data) were gridded with a geodesic grid in order to facilitate statistical comparisons on a global scale. All of the data points of a particular type falling within one grid cell were considered one occurrence, following the usage of Parrish et al.

(1982). For example, if 5 mosasaur records fell within a single geodesic grid cell, then those 5 records would be considered 1 mosasaur occurrence. Gridding and defining occurrences in this way de-emphasizes heavily sampled areas (e.g.. North America and

Europe in this study) (Parrish 1982).

A 9-frequency (9-v) geodesic grid was imposed on the point data (reptile and lithologic localities). The geodesic grid system was used rather than an orthogonal

(latitude-longitude) grid system in order to reduce grid-cell size and shape biases, which are particularly noticeable when data occur near the poles (Moore 1998). The 9-v frequency grid has about the same resolution as a 5° x 5° orthogonal grid, and has the advantage of nearly equal grid cell sizes and shapes. The relatively coarse 9-v resolution is appropriate for this study because of the uncertainty in paleogeographic reconstructions and in the placement of data with the lowest spatial resolution, where error in position was between 1° and 5° latitude or longitude. Related to the uncertainty in paleogeographic reconstructions is the tectonic displacement of sediments and fossils after their deposition (Moore 1998). 175

Rotational minimization was used to find the minimum number of occurrences for all of the point data of a particular age. Minimization is necessary because in the geodesic grid system, 5 or 6 grid cells can share one point. Six closely spaced data points could thus be gridded as 6 occurrences rather than 1 occurrence (Moore 1998).

Rotational minimization involves incrementally rotating and regridding data relative to the 9-v grid. The increment with the fewest number of occurrences is used for defining occurrences (see Moore, 1998, for details; data were minimized using software written by

T.L. Moore).

Data were minimized using their paleogeographic coordinates, and all of the point data of a particular age (reptile and lithologic data) were rotated together. Data were rotated in 1° increments, using an Euler pole of latitude 90° and longitude 0°. The paleogeographic coordinates were obtained using rotation calculations fi-om PGIS

Mac™.

Geographic Analysis of Gridded Data

The gridded data were plotted on the paleogeographic maps for geographic analysis using Maplnfo Professional®. The number of marine reptile fossil occurrences falling into predicted upwelling zones was obtained by querying their intersection (i.e., where the gird cell centroids intersected the predicted upwelling regions). In order to provide room for plotting errors, uncertainty in the paleogeographic reconstructions, and reptile mobility, I also queried the intersection of any part of the grid cell polygon with the predicted upwelling regions. The sides of the geodesic grid cells that represent occurrences are about 750 km long and are of nearly equal size no matter where they plot 176

on the globe. Allowing any part of the grid cell to intersect predicted upwelling creates a

buffer zone.

Some reptile fossils would be expected to fall into modeled upwelling zones by

chance. The percentage of shelf area occupied by predicted upwelling zones was

multiplied by the number of reptile occurrences to estimate the number of reptiles that

would occur in upwelling zones but would not necessarily be associated with upwelling

(following Parrish and Curtis 1982). The number of reptiles actually falling into

upwelling zones can then be compared with the number of occurrences expected in a

random distribution. Individual probabilities were calculated for the number of

occurrences falling within upwelling for each stage. The G-test for goodness of fit was

used to test the null hypothesis that reptile occurrences were distributed randomly across

the shelf during the entire Mesozoic (i.e., all data lumped together). This test is valid assuming that upwelling was equally important across all of the stages (Parrish and Curtis

1982). The percentage of shelf area occupied by upwelling ranged from 19.7 - 28.8 % over the Mesozoic (Table 3.3), so predicted upwelling assumed similar importance among intervals.

Distance Analvsis of Gridded Data

If marine reptiles were associated with upwelling, then it might be expected that marine reptile fossils and upwelling-related lithologies would be associated depositionally. The number of marine reptile fossil occurrences associated with upwelling-related lithologies was obtained by calculating the angular (arc) distance from 177

one fossil occurrence to all lithology occurrences for a given map interval. Equation 2

gives the angular distance relative to a unit sphere (Butler 1992):

ab = cos"' (sin Xa sin Ab + cos Aa cos At cos [a - <^b]), (2)

where ab is the distance between 2 points a and 6, Xg is the latitude of point a, and is

the longitude of point a. Equation 2 was modified to give the distance on the surface of

the earth;

ab = (cos"' (sin Xa sin Xb + cos Xa cos Xb cos [^a - ^6]))40076, (3)

where 40076 is the circumference of the earth in kilometers. If the shortest distance from a fossil occurrence to an upwelling-related lithology occurrence for a map interval is consistently small, then an association between them is suggested.

The distance from each reptile occurrence to each lithology occurrence was calculated using Equation 3. The shortest distance was obtained by identifying, for each reptile occurrence, the minimum distance calculated. For example, say that one calculated these distances from one mosasaur fossil occurrence to five lithology occurrences: 0 km, 550 km, 550 km, 1400 km, and 7200 km. The shortest distance between that mosasaur occurrence and any lithology occurrence is 0 kilometers. Only one shortest distance was used for each reptile occurrence. Thus, if a reptile occurrence was 550 kilometers away from several lithology occurrences (and 550 km was the 178 minimum distance calculated for that reptile occurrence), only one of those 550-kilometer distances was used in constructing the shortest distance histograms (note, however, that because gridded data were used, only 1 lithology occurrence could be 0 km away from a reptile occurrence).

It was possible to determine the range of distances represented by adjacent grid cells because gridded data were used, where distances were measured between cell centroids rather than between raw point localities (Figure J.1). For example, the

Cenomanian map interval contained the most grid cells, and the distances between cells of this stage were used to determine which grid cells were adjacent (sharing one side.

Case A of Table J.l). A distance of 550 km was used as the maximum distance between adjacent cells sharing one side.

The distributions of shortest distances were compared for both age-restricted and

-unrestricted data and the number of reptile and lithology occurrences that coincided or were one cell adjacent was counted. If reptile and lithology localities were closely associated, then one would expect the shortest distances to cluster in the 0-550 kilometer portion of the histogram (with 550 kilometers representing the operational upper limit of an occurrence within one grid cell adjacent, as explained above)—the distribution would be right-skewed. On the other hand, if there was not a close spatial association of reptile and lithology occurrences, one would expect the distribution of shortest distances to be either normal (ambiguous association) or left-skewed (negative association). A bimodal distribution with one mode between 0-550 kilometers and another mode at a greater distance would indicate that some reptile occurrences were closely associated with 179 upwelling lithologies whereas others were not. A multimodal distribution would be interpreted similarly. 180

RESULTS

Database and Data Processing

A total of 817 locality records comprising 1365 taxon-localities were entered in

the marine reptile database; five of these localities contained taxa that have been

associated with fresh water (5 taxon-localities total, see database methods and Tables

G.1-G.2). The number of localities and taxa increases from the beginning to the end of

the Mesozoic at all age resolutions, although diversity varies from interval to interval

(Figure 3.1, Table 3.4). The variability in diversity, as well as the overall increase over

time, are most likely related to preservation and effects of grouping the data (see

discussion). Grouping the data into map intervals introduces apparent declines in

localities for the Callovian and Maastricthian (however, sample sizes for the stage-

restricted data are smaller than for the unrestricted data) (Figures 3.1 and 3.2, Tables 3.4

and 3.5).

The number of upwelling-associated lithologies also increases over time, again

probably because of preservational and data-grouping effects (Figure 3.2, Table 3.5).

Both the marine reptile and the lithologic data were tightly clustered, as reflected by the decrease in number of occurrences after gridding and rotational minimization (Table 3.6, compare with Table 3.4). Clustering is particularly evident in the Cretaceous, where

numerous specimens have been recovered from sediments in North America (Figures I.l-

I.IO). The uncertainty in placement of western North American terranes is reflected in the position of several data points off the western North American shelf. For some map intervals, a few data points appear to plot on land but actually represent inland seas that 181 are not shown on the particular time slice of the map. This is particularly true for western

North America and Australia during the Cretaceous.

Geographic Analysis of Gridded Data

Whereas few marine reptile occurrences fell wholly within predicted upwelling, many more occurrences partially intersected predicted upwelling areas (Table 3.7,

"falling wholly within" defined as the grid cell centroid intersecting the upwelling area).

For about two-thirds of the map intervals, the number of marine reptile grid cells falling wholly within predicted upwelling was less than the number of grid cells predicted to fall within predicted upwelling by chance (Table 3.7). This pattern was observed whether all the data were included (series-level), the data were restricted to map interval stage, or the data were summed for the Mesozoic. On the other hand, about 80 % of the map intervals had more grid cells partially intersecting upwelling than would be expected by chance

(assuming that ideally, occurrences that partially intersect upwelling would have fallen wholly within the modeled areas). Exceptions occurred for the Valanginian (restricted and unrestricted data), Pliensbachian (restricted), and Anisian (unrestricted). At least half of all reptile occurrences partially intersected predicted upwelling for the following stages: Coniacian (restricted and unrestricted ages), Cenomanian (restricted and unrestricted ages), and Norian (restricted and unrestricted ages).

When considered together, the number of reptile occurrences that partially or wholly intersect predicted upwelling is statistically significantly greater than would be expected by chance {P - 0.05) for the following map intervals: Maastrichtian (restricted and unrestricted), Coniacian (unrestricted), Cenomanian (unrestricted), Aptian 182

(unrestricted), Caliovian (unrestricted), and Norian (unrestricted) (probabilities calculated directly. Table 3.7). The number of occurrences that are possibly explainable by upwelling (i.e., fall within or partially intersect predicted upwelling) is also statistically significant for the Mesozoic (all data lumped together) with both restricted and unrestricted data (G-test, Gadj = 4.7144 for restricted, Gai/y=6.8438 for unrestricted, P

= 0.05 in both cases. Table 3.7). For all of the data together, the proportion of reptile occurrences partially or wholly intersecting upwelling was virtually the same for age- unrestricted as for age-restricted data (unrestricted, 43% and 16% respectively; restricted,

44% and 15% respectively).

All marine reptile families in the database had occurrences that intersected predicted upwelling. Families in which half or more occurrences intersected predicted upwelling included the Shastasauridae (Ichthyosauria); the Elasmosauridae and

Polycotylidae (Plesiosauria); and the Mosasauridae (particularly Tylosaurinae and

Plioplatecarpinae). Many specimens unidentified to the family level for all three orders also intersected predicted upwelling (Table 3.8). The following map intervals had more than half of the occurrences for a majority of reptile families falling into the upwelling buffer; Maastrichtian, Coniacian (restricted and unrestricted), Cenomanian (restricted and uiu-estricted), Aptian (unrestricted), Caliovian (restricted and unrestricted), and

Norian (unrestricted). These are most of the same intervals that tested statistically significantly from a random distribution at /* = 0.05 (see above and Table 3.7).

For age-restricted data, marine reptile taxa with more occurrences intersecting upwelling than would be predicted by chance include the Plesiosauria and Mosasauridae 183 as a whole and the Pliosauridae, Polycotylidae, Mosaurinae, Plioplatecarpinae, and

Tylosaurinae (Table 3.9, P = 0.05). The same relationships hold for age-unrestricted data, with the addition of the Ichthyosauria as a whole and the Shastasauridae. The number of reptile occurrences in the Ichthyosauria, Plesiosauria, and Mosasauridae indeterminate categories also intersected upwelling more than was predicted by chance

(Table 3.9, P = 0.05).

Distance Analysis of Gridded Data

Shortest distances were a function of the number of reptile localities, and the number of shortest distances per time interval ranged from 3-63 (Table 3.10). Comparing the histograms for each map interval (Figures J.2-J.11), right skewness became more pronounced as sample size increased. The histogram for each time interval had a mode in the 0-kilometer bin except for the Anisian (age-unres.) and Pliensbachian (age-res.), both of which had observations in the 0-kilometer bin. Histograms for the Maastrichtian,

Coniacian (except age-res.), Cenomanian, Valanginian (very small sample) and

Kimmeridgian showed definite right-skew. Histograms for the Aptian, Callovian,

Pliensbachian, and Norian had a 0-kilometer mode and right-skew, but also had clusters of observations in the right tail (longer distances). The histogram for the Anisian was centered on distances greater than those expected for adjacent grid cells (>550 km) and showed the least amount of right skew of all of the histograms.

Not surprisingly, the mean, minimum, and maximum shortest distances between reptile and lithology occurrences usually were smaller for age-restricted than for age- unrestricted data (Table 3.10). For age-restricted data, mean shortest distances for all 184

lithology types together were less than 550 kilometers (1 cell adjacent, sharing one side)

for the following map intervals: Maastrichtian, Coniacian, Cenomanian, Valanginian,

and Pliensbachian. Interestingly, no reptile occurrences intersected predicted upwelling

for the Valanginian and Pliensbachian map intervals (Table 3.7), yet reptile and lithology

occurrences were close together in these time periods (Table 3.10; note the small sample

sizes).

No particular lithology type dominated the average shortest distances (Table

3.10), but different lithology types were close to reptile occurrences at different times.

For the Pliensbachian interval, reptile occurrences were on average closest to organic-rich

rock, phosphorite, and glauconite. In the Valanginian, reptile occurrences were on

average closest to phosphorite. Throughout the Cretaceous, reptile occurrences were on

average closest to glauconite and to grid cells with more than one lithology present

(Table 3.10, Appendix I, "c" maps).

Similarly, no particular reptile order (or family, in the case of Mosasauridae) was

consistently closer to upwelling-related lithology occurrences than other orders (Table

3.11). Again, average distances for age-restricted data tended to be smaller than for age-

unrestricted data. Pliensbachian, Kimmeridgian, and Cretaceous map intervals tended to

have the shortest average distances for age-restricted data. The Early Cretaceous

intervals had very small sample sizes that contributed to the "0" averages (each reptile occurrence coincided with a lithology occurrence).

The ratio of grid cells that contained both reptile fossils and upwelling-related lithologics varied from 25 - 71 % of age-restricted reptile occurrences (except for the 185 three oldest map intervals, there were more lithology than reptile occurrences) (Table

3.12). When adjacent cells that share one side (as opposed to a vertex, see Figure J.l) were included, the ratio varied from 40-100 %. No particular lithology type dominated these cells, although from the Middle Jurassic on, a majority of cells tended to have more than one upwelling-related lithology present (Table 3.13). No single reptile taxon dominated the cells, but over the Mesozoic, certain higher taxa made up a majority of cells: ichthyosaurs in the Triassic, ichthyosaurs and plesiosaurs in the Jurassic, plesiosaurs in the Early Cretaceous, and mosasaurs in the Late Cretaceous (Table 3.13). 186

DISCUSSION

For the Mesozoic data together, marine reptile occurrences are distributed non- randomly (statistically) relative to predicted upwelling areas. At the stage level with age- unrestricted data, the correspondence between reptile occurrences and predicted upwelling tested nonrandom for all four of the middle and Upper Cretaceous map intervals but for only two older intervals (Callovian and Norian). These results are based on the partial intersection of grid cells; relatively few grid cells fell wholly within predicted upwelling, but that is probably related to the narrow shape of predicted upwelling areas and the relatively large size of the grid cells. These results suggest that upwelling could have played a role in marine reptile distribution, particularly in the latter half of the Cretaceous.

Marine reptile occurrences also fell outside of predicted upwelling, and some of the occurrences that fell within upwelling might not have been related directly to upwelling. The spatial distribution of marine reptile fossils for a given map interval does not appear to be simply random, however. The statistical tests described above suggest a nonrandom distribution relative to upwelling area for the Mesozoic together and for six map intervals (Table 3.7). Also, visual inspection of the data shows that the fossil localities appear to be clustered, particularly near inland seas and modeled upwelling areas (Figiires 1.1-1.10). Sampling, preservational, and analytical biases, and particularly other aspects of marine reptile ecology, more likely influenced the results. Discussion of each of these issues follows. 187

Sampling Biases

It is more difficult to obtain a statistically significant probability as sample size

decreases; n must equal 3 or more in order to obtain a significant probability using

binomial probabilities (n is at least 3 for all map intervals in this study; see Table 3.7).

For small sample sizes, the null hypothesis (random distribution across the shelf) might

not be rejected even if the occurrences are actually non-randomly distributed. Map

intervals with more occurrences on the shelf provide more robust tests that the

occurrences are randomly distributed. Several factors affect the sample size for each

map interval, including search effort and monographic effects.

The statistics used to calculate probabilities for fossil localities corresponding

with predicting upwelling areas assume random sampling of fossil localities over the

entire shelf, in other words, that all of the published reports in the literature have sampled

the shelf randomly and completely. This assumption is not met because, first of all, not

all of the shelf for a given geologic period is available for sampling. Second,

paleontologists tend to search for fossils in and around established localities, so that the

more fossils found in an area, the more that area is searched. Finally, more fossils are

found where an area has been searched for a long time (Sander et al. 1997). In order to

compensate for the effects of search effort, a concerted effort was made to seek out

literature from areas such as Russia and northern South America that might not have been

searched as intensely as North America and Europe. The bias towards Europe and North

America is particularly noticeable for the Late Cretaceous (Figures 1.8 -1.10), but inland seas covered those areas as well. If reptiles were known only from those two continents 188 in the Late Cretaceous, then the search effort bias would be strong. However, marine reptile fossils are also known trom Africa, South America, China, and Antarctica during those times. Marine reptile fossils are known from most continents throughout the

Mesozoic (Figures I.l-I.lO). Search effort is probably not the main factor contributing to the fossil distribution.

Marine reptile fossils are possibly under-reported if an area has been searched for only a particular kind of fossil or if only the well-preserved specimens are published.

Welles and Gregg (1971, p. 101) noted that "the literature does not reflect the total fauna as investigators usually have been concerned with a single specimen or the larger ones."

On the other extreme, bone fragments found by geologists with little background in paleontology could result in isolated occurrences being under-reported. Hidden specimens—unidentified, unreported material stored in museum drawers—also contribute to the under-reporting of marine reptile distribution. Under-reporting can be partly compensated for by conducting a broad literature search not restricted to reports focused on vertebrate paleontology. A survey of the database fields relating to taphonomy shows that a variety of marine reptile finds are included, not just exceptional specimens (Table 3.9). Thus, a strong bias towards exceptional specimens is not present in this study.

Sampling is potentially altered by monographic effects. Different species concepts employed by taxonomists studying the same order or family over a different time interval can result in spurious diversity estimates. This study uses higher taxonomic levels (orders and families) for the analysis, where monographic effects are not as 189 pronounced (Raup 1976). Spatial data distribution is also more important for this study than temporal trends of diversity at lower taxonomic levels. Another potential monographic effect characteristic of older literature is the regional naming effect. Fossils with similar morphologies and of the same clade and age but from widely separated localities (usually different continents) are sometimes identified as separate species.

Revision of these specimens often results in lumping of taxa (e.g., Bardet 1992). The regional naming effect was addressed by consulting more recent literature for current taxonomic schemes (Tables E.I-E.3), with the result that taxa that were regionally divided were often lumped (e.g., some specimens of Ancanamunia from South America can be synonymized with Ophthalmosaurus, a cosmopolitan genus). Because this study uses families and orders for analysis, monographic effects are minimal and do not alter the overall spatial relationship between predicted upwelling and marine reptile fossils.

Preservational Biases

Underlying literature-based sampling biases are biases related to the amount of rock exposed for searching. A larger sample size for a map interval increases the chances that a given locality will be associated with upwelling. The Coniacian interval, for example, contains the most reptile occurrences and the most occurrences possibly explainable by upwelling (Table 3.7). The number of localities for a time period will be related to the amount of sedimentary rock preserved, exposed, and accessible for that age.

For example, Raup (1976) showed that Phanerozoic diversity and the area of exposed sedimentary rock follow very similar trends. Times of relatively higher sea level mean that potentially more continental area is flooded and potentially more shelf sediment is 190

preserved. In this study, the first-order increase over time in number of fossil and

lithologic data points follows the same trend as sea-level curves published for the

Mesozoic (Hallam 1992). Large areas of Cretaceous sedimentary rock are exposed (see

Figure 2 in Raup 1976), and the largest numbers of vertebrate localities in terms of both

raw and gridded data occur in the Cretaceous (Figure 3.1, Tables 3.4 and 3.6). Jurassic

localities and occurrences are more numerous than Triassic localities, despite the fact that

Jurassic sedimentary rocks occupy less area today than Triassic rocks (see Figure 2 in

Raup 1976). This result could reflect better accessibility to or more thorough searching

of Jurassic than Triassic sediments.

This study is necessarily restricted to shelf environments because most of the

deep-sea record has been destroyed through subduction. Any marine reptiles that were

buried in deep-sea sediments will be under-sampled. No well-documented, deep-sea

marine reptile fossils are currently known (Martill et al. 1995), although they are

potentially preservable given that whale bones and whole whale carcasses occur today on

the deep-ocean floor (Eastman 1903; Allison et al. 1991; Martill et al. 1991). The database employed in this study contains only two possible deep-ocean localities (NA-

682, NA-683, Franciscan Group, Tables G.l and 0.2), and their age and the lower-level taxonomy of the fossils are uncertain. The relative proportion of coastal versus pelagic dwelling marine vertebrates will affect the ratio of carcasses deposited in shallow- versus deep ocean-sediments. Today, pelagic baleen whales have a greater chance of being buried in deep sediments than neritic cetaceans such as gray whales, who spend their 191 lifetime on the shelves (Rice and Wolman 1971). Potential preservational differences between deep-sea and shelf environments are beyond the scope of this study.

Marine reptile fossils could potentially be better preserved in upwelling areas, regardless of their feeding strategies, simply because of the high sedimentation rates and more extensive oxygen minimum zones that characterize these areas (see Chapter 1). For example, a predicted upwelling area could occur in a region where current-drifted animals finally sink or get washed ashore (see, e.g., Schafer 1972, for today's North Sea).

Detailed information on fossil preservation and local paleogeography and sedimentology is necessary to determine the role of current-drift in a particular fossil deposit. Deposits consisting of current-drifted specimens can potentially represent spatially averaged assemblages (Barnes 1977). The results of this study show that a preservational bias in predicted upwelling areas is probably not present, because as many reptile occurrences are located outside predicted upwelling as inside (Table 3.7). Exceptional preservation is also not limited to predicted upwelling areas (Table 3.9).

The nature of fossil locality data illustrates another potential preservational bias, that of patchy fossil distribution. McGowan (1978) noted that the ichthyosaur genera

Ophthalmosaurus, Ichthyosaurus, Mxosaurus, Platypterygius are widely but spottily distributed. Considering only localized exposures of a taxon can give the impression of a restricted geographic range. Additionally, the patchy nature of fossil distribution affects search effort, leading to oversight of less abundant or more scattered fossil localities.

Gridding and performing rotational minimization on the data reduced the effects of oversampling rich fossil localities, while retaining information on isolated localities. The 192

gridding techniques, coupled with a global-scale database, can help assess whether a

patchy fossil distribution results mainly from preservation or from the animals' ecology.

Analytical Biases

Details of the data can be obscured in a study such as this one where localities are

grouped into map intervals and where higher taxonomic levels are used for analysis.

Grouping the data into map intervals means that some localities might be plotted on the

"wrong" stage. This is particularly evident for sea-level changes that affect inland seas.

The apparent decline of localities and taxon-localities for the Maastricthian probably

results from placing Upper Cretaceous localities in the Coniacian map interval (except for

Upper Cretaceous ichthyosaur localities, which were placed in the Cenomanian interval,

see Table 3.2.b). Data were generally placed in a map interval that fell as close to the

middle of the epoch as possible (Table 3.2). Using data subsets for time-restricted and

-unrestricted localities provided a way to assess the effects of data grouping. Data

grouping appears to have enhanced results for occurrences possibly explainable by

upwelling for the Coniacian, Aptian, Kimmeridgian, and Norian. Results for the

Maastrichtian, Cenomanian, and Callovian more likely reflect actual patterns because

they tested statistically significant for the both the stage-restricted and stage-unrestricted

cases (Table 3.7).

The upwelling model might not accurately reflect the distribution of ancient

upwelling. Between one- and two-thirds of the upweliing-related lithologies intersected

predicted upwelling (Table 3.10), with higher correspondences in the middle and Late

Cretaceous intervals. Higher correspondences were also observed for occurrences with 193 multiple lithology types present than for single lithology occurrences. Phosphorite, glauconite, and organic carbon had higher correspondences than biogenic silica. Even in modem upwelling areas, though, not all upwelling-related lithologies are present (see

Chapter 1), and a combination of lithology types is often a better indicator than a single type.

Marine reptiles were relatively close to upwelling-related lithologies for time periods when their occurrences did not intersect upwelling significantly (e.g.,

Pliensbachian, Valanginian). The upwelling predictions or the sedimentary interpretations could be in error; the variation in sample size for the angular distance calculations also affects the results (Tables J.l-J.lO). Looking at the Pliensbachian maps, though (Figure 1.3), shows that lithologies are clustered near sites of predicted upwelling even though they do not intersect the areas, implying that the lack of correspondence could result from paleogeographic uncertainty or plotting errors. The number of associated reptile and lithology occurrences increases by 30-70% when one grid cell adjacent (side shared) is included, again suggesting clustering of lithologies and reptile fossils (Table 3.12).

Lithologies tend to cluster near predicted upwelling. Exceptions include isolated occurrences of glauconites, which can be common in nearshore environments whether or not upwelling occurs, and biogenic silica loceilities on the western sides of ocean basins

(particularly the western Pacific), which could reflect dynamic upwelling associated with western boundary currents. Reduced correspondence between the lithological and model comparisons might also resxilt firom modeling only certain types of upwelling (mainly 194

coastal and high-latitude divergences on the shelf that are easier to infer for the past,

Parrish 1982; Parrish and Curtis 1982).

Comparing reptile fossil distribution with predicted upwelling distribution

introduces a possible bias because the actual proportion of upwelling to non-upwelling

sediments exposed (and available for searching for marine reptiles) is unknown. The

proportion of upwelling to non-upwelling sediments preserved and exposed might differ

from the proportion predicted by the model. A difference would affect the statistical

calculations, which are based on shelf area calculated from the paleogeographic

reconstructions and model predictions, not on the area of shelf sediment exposed. If

much more upwelling sediments are exposed than non-upwelling sediments, the

proportion will be larger than the model predicted, more reptiles will be expected to fall

into predicted upwelling by chance (simply because more upwelling sediments are

exposed for searching), and reptile distribution will be predicted by rock exposed.

Determining rock exposed for any time interval is a large undertaking, and uncertainty

remains because one cannot know how much rock is missing. Upwelling-related

lithologies tend to cluster in certain areas of the shelf (Appendix I) and it seems unlikely

that a considerably larger area of these lithologies would be preserved than other shelf sediments. The modeled upwelling predictions are the best estimates available at this

time.

Even though proportionally more ancient shelf area is preserved than deep-ocean area, much is missing. Where sedimentary data are lacking, locations and sizes of continental shelves are extrapolated on the paleogeographic reconstructions. The effects 195

of missing shelf mean that area calculations are very rough estimates, and changes in

shelf area would affect the statistical analyses because upwelling area is partially a

function of the shelf area. If predicted upwelling areas are underestimated, fewer

occurrences would be expected to fall into predicted upwelling by chance (lower

percentage of upwelling area = fewer occurrences expected to randomly fall into

upwelling). This would make it easier to get a significant result that the distribution is

non-random (vice versa for the case if predicted upwelling area is overestimated). Area

estimates were similar to those of Parrish (1985) for shelf and Parrish and Curtis (1982)

for upwelling areas (shelf areas differ from 4-20 %; upwelling areas from 2-5%, Table

3.3). The differences result from using different reconstructions and methods (Parrish,

1985, used a planimeter rather than a grid to calculate area). The overall similarity

between results suggests that the areas used here are reasonable estimates.

Marine Reptile Ecology

Of the marine reptile families sampled, at least half of the occurrences containing

members of the Shastasauridae (Ichthyosauria); Elasmosauridae and Polycotylidae

(Plesiosauria); and Tylosaurinae were found within the predicted upwelling buffer (Table

3.8). All of these families include species that attained large sizes (> 2 meters long).

Most members of the Pliosauridae and Tylosaurinae were top predators with teeth equipped for taking large prey, while members of the ichthyosaur families had teeth capable of taking soft prey such as unshelled or internally shelled cephalopods (Massare

1987). The elasmosaurids and polycotylids were probably generalist feeders (Massare

1987); elasmosaurs could perhaps feed by darting their long necks in and out of schools 196 of prey such as fish. Based on ichthyosaur distribution in the former Soviet Union,

Nesov (1988) suggested that ichthyosaurs, especially in the later part of their existence, lived in upwelling zones. Large animals and top predators require large amounts of food, and high food requirements can be met where smaller marine animals congregate, including upwelling zones and shallow coasts along migration routes, breeding grounds, or estuaries (Slijper 1979; Wiirsig 1989).

Aside from the three families discussed above, it appears that age was a more important factor in whether or not a taxon fell within the predicted upwelling buffer. For certain time periods (Maastrichtian, Coniacian (unrestricted), Cenomanian (restricted and unrestricted), Aptian (unrestricted), Callovian (restricted and unrestricted), and Norian

(restricted and unrestricted)), half or more of the reptile occurrences fell within the predicted upwelling buffer, regardless of the taxon (Table 3.8). The Cretaceous intervals were times of high relative sea level, providing more shelf habitat and a greater absolute

(though probably not relative. Table 3.3) area for upwelling and productive shelf habitat.

Marine reptile diversity was particularly high in the Mid-Late Triassic and in the Late

Cretaceous (Bardet 1994). Collin and Janis (1997) noted that these times of diversification corresponded to the times when organic-rich rocks were statistically significantly correlated with predicted upwelling as shown by Parrish and Curtis (1982).

A plentiful marine food supply could have triggered diversification of marine reptiles

(Collin and Janis 1997). Multiple radiations into the high-productivity, top predator niche are suggested by the predominance of different higher taxa in shared reptile- upwelling-related-lithology occurrences over the course of the Mesozoic: ichthyosaurs in 197 the early Mesozoic, plesiosaurs in the middle Mesozoic, and mosasaurs in the late

Mesozoic (Table 3.13).

Mobility, reproductive behavior, latitudinal restrictions, and stranding are factors besides feeding ecology that could potentially affect marine reptile distribution. Reptile mobility increases the area over which a dead animal could fall and be buried. Many baleen whales migrate between feeding grounds, often located in cold, food-dense, waters, and calving grounds located in warmer, sheltered waters (Gaskin 1976; Gaskin

1982). Gray whales today feed in the productive, shallow Bering Sea, migrate down the western North American coast, and give birth in lagoons off the Baja California coast

(Rice and Wolman 1971). Note that although gray whales migrate through shelf affected by upwelling (the Oregon, California, and Baja shelves), they do not feed heavily in these areas. Therefore, a paleontologist working in the future might find gray whale skeletons in coastal or shallow shelf deposits associated with California Current upwelling, but the association resulted from migration, not a reliance on the upwelling area for feeding.

Assuming that some marine reptile taxa had similar behaviors, we could expect migration to similarly influence the distribution of reptile carcasses. Elderly or diseased animals might be expected to die during migration, but without knowing details of the animal's migrating behavior, no unique characteristics could distinguish a death along a migratory path from a death in a feeding area.

Another implication of animal mobility is that opportunistic animals are likely to range widely in search of food. These animals could forage in upwelling areas and in productive shallow shelves. Shallow inland seas have been suggested as sites of high 198

shelf production, whether by upwelling (Parrish and Gautier 1993), shelf recycling, or

high nutrient runoff that would promote algal blooms (Hudson and Martill 1991).

Opportunistic feeders could take advantage of the resources available in upwelling areas

but not necessarily depend on them.

Some whale taxa (notably baleen whales) spend the winter in low-latitude,

relatively unproductive waters for the calving season (see Whitehead and Moore 1982,

for humpback whales in the West Indies). Birthing or egg-laying areas could have

formed sources of marine reptile fossils that would be potentially uru-elated to upwelling.

Very young individuals would likely die only in or very near the birthing ground; adults

could be stranded after egg-laying. Dead reptiles would be buried within the same basin

and perhaps even in the birthing ground (e.g., if a lagoon), depending on the fat content

of the carcass, water depth, and current patterns within the basin (Schafer 1972). These

factors all affect the condition of carcass upon burial and how quickly it is buried. The

main criterion that paleontologists use for interpreting a marine reptile deposit as a

breeding ground is the presence of very small individuals that are interpreted as embryos.

Possible embryonic material of ichthyosaurs has been recovered from the Toarcian

Posidonia Shale in Germany, the Lower Jurassic of Somerset, England, the

Anisian/Ladinian Grenzbitumenzone in Switzerland, and the Camian/Norian Luning

Formation in Nevada (Massare 1988a; Martill 1995). All four deposits were located in

low to mid-latitudes in the more basinal parts of epicontinental seas or small epicontinental basins. The Posidonia Shale and the Grenzbitumenzone represent envirotmients of exceptional preservation (Martill 1993; Furrer 1995), so the relative 199

scarcity of reported embryonic material from other regions might reflect a bias against its

preservation (small, delicate bones).

Marine reptiles could have had temperature requirements that limited their

distribution to certain latitudes. However, the paleobiogeography of all three reptile

orders sampled in the database spans equatorial and high latitudes (Figures 1.1-10). At

least some members of these marine reptile orders do not appear to be restricted to low

latitudes.

Marine reptiles might have been prone to stranding, as whales are today.

Stranding occurs when whales get caught in a ebbing tide in very shallow waters or

otherwise become beached. This phenomenon causes the death of numerous cetaceans in

the North Sea (Schafer 1972). The North Sea appears to act as a sperm whale trap

because sperm whale strandings seem to occur during the autumn migration, when

individuals are weakened and more prone to getting caught in the shallow parts of the

basin (Jauniaux et al. 1998). Evidence for current-drifted and refloated specimens (see

preservational biases, above) and also for bones in coastal or beach sediments would support stranding as a factor affecting the marine reptile deposit.

In suirmiary, several ecological factors besides feeding ecology can affect marine

reptile distribution. However, one of the factors that is easiest to test in a large-scale comparison such as this one is feeding ecology, specifically in regard to upwelling. For the other factors, detailed taphonomic and sedimentologic information about each fossil locality is needed in order to determine their relative contribution. A locality-by-locality analysis of these additional factors is beyond the scope of this paper, but the results of 200 this study indicate that upwelling may well have influenced marine reptile distribution, particularly for the latter part of the Cretaceous Period. 201

CONCLUSIONS

Upwelling could have affected marine reptile distribution by providing feeding grounds. When partial grid cell intersections are allowed, marine reptile fossil localities are non-randomly distributed relative to predicted upwelling zones for six of the ten map intervals used in this study. Four of these intervals are in the middle-Late Cretaceous.

During these times, relatively more shelf area was available for upwelling and shelf production to occur. In some time intervals marine reptile localities were close to upwelling-related lithologies and did not intersect predicted upwelling, but plotted nearby.

Marine reptile families with half or more of their occurrences consistently falling into predicted upwelling include taxa with diverse feeding habits such as large, killer whale-type predators, generalist feeders, and soft-bodied prey feeders. The families all contain taxa that tend towards large body size, including occurrences of three of the four largest known marine reptile taxa mentioned in the introduction. A large body size would necessitate an abundant food supply, which could be found in upwelling areas

(though several other large-bodied taxa were not associated as much with predicted upwelling). Several other aspects of marine reptile ecology likely affected their distribution, including reproductive and migratory behavior and stranding, and the relative contributions of these factors can be discovered through site-by-site taphonomic and sedimentologic study.

Five of the six time intervals where reptile occxirrences tested non-random relative to predicted upwelling have been identified as times of peak diversification for marine 202 reptiles in general (Middle-Late Triassic and Late Cretaceous; Bardet 1994). During the

Mesozoic, at least 16 lines of diapsid reptiles radiated into the oceans (Carroll 1985), and a food-rich environment might encourage such radiation (McGowan 1972; Collin and

Janis 1997). Conversely, a decline in production could adversely affect marine reptile diversity and abundance. The mosasaurs radiated rapidly in the Late Cretaceous and died out just as suddenly at the end of the Cretaceous. The Mosasauridae contained more occurrences that intersected predicted upwelling than any other reptile family (Table 3.9).

Mosasaur fossils are conomon in shallow-water, chalky and/or phosphatic sediments, suggesting high productivity. Most taxa ate large prey and probably were slow- swimming ambush predators (Massare 1987; 1988b). Bardet (1994) suggested that a drop in primary production could have contributed to their demise, and a decline in marine productivity has been postulated for the Cretaceous-Tertiary boundary (Zachos et al. 1989; Coccioni and Galeotti 1994). The results of this study support the idea that primary production influenced Mesozoic marine reptile distribution and diversification. 203

FIGURE 3.1. Number of localities and taxon-localities for differing temporal resolutions. See Table 3.2.a. for stage codes. Compare with Figure 3.2, which shows all data grouped into the time periods used for gridding.

Stage-restricted data

• Families •Taxon-localities • Localities

50 100 150 200 250 300 350

h i

9 e

e & d Stage-unrestricted data

•Families b BHH •Taxon-localities • Localities a

50 100 150 200 250 300 350 204

FIGURE 3.1. (continued)

Series-unrestricted data

• Families •Taxon-localities • Localities

0 100 200 300 400 500 600 700 205

FIGURE 3.2. Number of marine reptile and lithologies localities by map interval. Numbers represent age-unrestricted, ungridded point data, a) all reptile and lithologies localities, b) lithology types, c) higher-level reptile clades (next page)

A.

Maa

Con

Cen

Apt

Val

Kim

Cal • Lithologies

PI! • Reptile localities

Nor El Reptile taxon- An! localities

100 200 300 400

B.

•Organic-rich rock • Phosphorite • Biogenic silica BGIauconite

50 100 150 200 250 300 350 400 206

FIGURE 3.2. (continued)

c.

• Mosasauridae • Plesiosauria • Ichthyosauria

Ani ||u

50 100 150 200 250 300 350 400

Map interval codes: Ani = Anisian, Nor = Norian, Pli = Pliensbachian, Cal = Caliovian, Kim = Kimmeridgian, Val = Valanginian, Apt = Aptian, Cen = Cenomanian, Con = Coniacian, Maa = Maastricthian 207

TABLE 3.1. Stages of paleogeographic maps used in data plotting and analysis. Maps were obtained from PGIS Mac™.

Stage Age of Reconstruction, Ma Maastrichtian 69.4 Coniacian 88.0 Cenomanian 94.0 Aptian 118.0 Valanginian 130.2 Kimmeridgian 152.2 Callovian 166.0 Pliensbachian 195.0 Norian 216.0 Anisian 237.0 208

TABLE 3.2.0. Codes used to assign data points to a map interval for plotting. Numerical ages from Harland et al. (1990). Stages shown in capitals denote map intervals. Grouped stages duration refers to the duration of intervals in the stage_un category (see Table 3.2.b). Stage_res = restricted stage, Stage_un = unrestricted stage, and series un = series. Ma Duration, Ma Stage_res, Lower Upper Grouped Period Epoch Series un Stage Stage_un Bound Bound Stage Stages Series Cretaceous Upper h MAASTRICHTIAN j 74.0 65.0 9.0 9.0 32.0 Campanian 83.0 74.0 9.0 Santonian 86.6 83.0 3.6 CONIACIAN i 88.5 86.6 1.9 16.4 Turonian 90.4 88.5 1.9 CENOMANIAN h 97.0 90.4 6.6 Lower g Albian 112.0 97.0 15.0 21.6 48.6 APTIAN 8 124.5 112.0 12.5 Barremian 131.8 124.5 7.3 19.8 Hauterivian 135.0 131.8 3.2 VALANGINIAN f 140.7 135.0 5.7 13.8 Berriasian 145.6 140.7 4.9 Jurassic Upper f Tithonian e 152.1 145.6 6.5 11.5 KIMMERIDCIAN 154.7 152.1 2.6 11.5 Oxfordian 157.1 154.7 2.4 Middle e CALLOVIAN d 161.3 157.1 4.2 20.9 Bathonian 166.1 161.3 4.8 Bajocian 173.5 166.1 7.4 20.9 Aalenian 178.0 173.5 4.5 Lower d Toarcian 187.0 178.0 9.0 30.0 PLIENSBACHIAN c 194.5 187.0 7.5 Sinemurian 203.5 194.5 9.0 30.0 Hettangian 208.0 203.5 4.5 Triassic Upper c Rhaetian 209.5 208.0 1.5 27.0 NORIAN b 223.4 209.5 13.9 15.4 Camian 235.0 223.4 11.6 Middle b Ladinian 239.5 235.0 4.5 6.1 ANISIAN a 241.1 239.5 1.6 21.6 Lower a Spathian 241.9 241.1 0.8 3.9 Nammalian 243.4 241.9 1.5 Induan 245.0 243.4 1.6 209

TABLE 3.2.b. Criteria for assigning data points to a map interval for plotting. Map interval codes are shown in Table 3.2.a. Map Interval Code Criteria Stage res Restricted Stage Included data reported to stage level for one of the ten mapped stages. Data with ages that included a map stage as an upper or lower boundary or that ranged through a map stage were not included. Data had to fall wholly within the map stage.

Values: a-j, $ signifies a null value (data not reported to restricted stage level)

Stage_un Unrestricted Stage Included all data reported to the stage level. Stages were grouped as evenly as possible into ten intervals (see Table 3.2.a. for interval durations). Data reported to stage level but ranging through several stages were included. If data ranged through more than one map interval, the data were assigned to the older mapped stage, unless the known taxonomic age range contradicted the assignment.

Values: a-j, $ signifies a null value (data not reported to restricted stage level)

Series_un Series Included all data. Data ranging through more than one series (e.g.. Upper Jurassic - Lower Cretaceous) were assigned to the older series, unless the known taxonomic range contradicted the assignment (for example, no mosasaurs are known fi-om sediments older than Turonian, so a mosasaur locality dated as "Upper Cretaceous" was coded: $ - stage_res $ - stage_un h - series_un, and plotted on the Coniacian map). Data of Upper Cretaceous age were plotted on the Coniacian map, and data of Lower Cretaceous age were plotted on the Aptian map.

Values: a-h TABLE 3.3. Area in km' of continental shelf and predicted upwelling areas for this study and from previous studies. In this study, area was calculated at a 2° resolution using the Paleocliniate parametric climate model. For comparison, area estimates are shown from Parrish, 1985, for continental shelf and from Parrish and Curtis, 1982, for % shelf area occupied by predicted upwelling.

Time Paleoclimate Paleoclimate Paleoclimate Parrish, 1985 Parrish & Curtis, 1982 Ma Shelf area Upwelling area % shelf area Shelf area % shelf area km^ km" occupied by upwelling km^ occupied by upwelling 69.4 5.97E+07 I.24E+07 20.76 5.58E+07 22.10 88.0 6.30E+07 I.47E+07 23.37 94.0 6.66E+07 1.61E+07 24.23 5.55E+07 26.08 118.0 7.61E+07 I.50E+07 19.73 130.2 4.86E+07 1.28E+07 26.31 152.2 5.20E+07 I.31E+07 25.13 5.12E+07 21.95 (Volgian) 166.0 5.20E+07 1.04E+07 19.93 195.0 4.43E+07 I.25E+07 28.12 3.56E+07 32.66 216.0 4.60E+07 1.24E+07 26.98 237.0 4.50E+07 1.30E+07 28.83 4.32E+07 26.3 (Induan) 211

TABLE 3.4. Number of localities, taxon-localities, and families for differing temporal resolutions. The number of specimens indeterminant as to family is also shown. See Table 3.2.a. for stage codes. # of Loc # of Taxon Loc # Families indet. incertae sedis Stage-restricted a 0 0 0 0 0 b 13 16 5 7 2 c 7 11 5 1 1 d 15 31 4 9 0 e 40 65 5 32 0 f 6 6 1 5 0 g 13 23 7 7 0 h 30 34 4 16 0 5 5 3 0 0 119 231 5 31 0 Stage-unrestricted a 66 102 4 31 15 b 24 37 8 24 3 c 81 151 7 38 3 d 46 82 6 22 1 e 99 196 7 94 0 f 19 21 5 7 0 g 16 27 7 7 0 h 69 86 4 33 0 i 187 298 7 18 0 j 119 231 5 31 0 Series-unrestricted LTr 14 18 2 10 6 MTr 47 76 3 20 10 UTr 44 66 8 37 3 U 96 177 7 58 3 MJ 49 85 6 22 1 UJ 105 202 7 99 0 LK 84 110 7 42 0 UK 378 631 8 78 0 212

TABLE 3.5. Number of marine reptile and upwelling-related lithologies by map interval. Numbers represent age-unrestricted, ungridded point data. Data type Map Intervalt Ani Nor Pli Cal Kim Val Apt Cen Con Maa Total ta.xon-localities 94 66 177 85 202 21 45 86 358 231 Total localities 61 44 96 49 105 19 33 69 222 119 Ichthyosauria indet. or incertae sedis 43 26 34 12 55 7 11 20 Ichthyosauridae 1 13 10 21 Leptopterygiidae 1 15 Mixosauridae 22 2 Platypterygidae 1 2 3 20 Shastasauridae, Shonisauridae 22 19 Stenopterygidae 38 1 Temnodontosauridac 14 Total Ichthyosauria 87 49 114 23 77 9 14 40 0 0 Plesiosauria indet. or incertae sedis 3 14 27 II 44 8 13 30 31 Cryptoclididae 9 19 1 Elasmosauridae 1 8 18 1 3 9 15 54 32 Pistosauridae 4 1 Plesiosauridae 1 15 2 3 2 4 I Pliosauridae 13 22 57 7 6 10 26 4 Polycotylidae 1 4 8 29 6 Total Plesiosauria 7 17 63 62 125 12 31 46 141 73 Squamata VIosasauridae indet. or incertae sedis 52 46 Mosasaurinae 80 64 Plioplatecarpinae 61 42 Tylosaurinae 24 6 Total Mosasauridae 217 158 Upwelling-related lithologies Glauconite 7 5 8 51 30 51 80 146 79 131 Phosphorite 6 6 8 5 9 15 2 34 15 40 Biogenic silica 3 39 19 30 79 38 34 94 55 65 Organic-rich rock 3 3 2 8 6 5 12 23 15 13 Total lithologies 19 53 37 94 124 109 128 297 164 249 1274 t map interval codes: Ani = Anisian, Nor = Norian, Pli = Pliensbachian, Cal = Callovian, Kim = Kimmeridgian, Val = Valanginian, Apt = Aptian, Cen = Cenomanian, Con = Coniacian, Maa = Maastricthian 213

TABLE 3.6. Rotational minimization results. Reptile and lithologic data were rotated together for each map interval. The coordinates for the best step results were used to define grid cells for mapping. Best step refers to the rotation angle which gave the fewest number of occurrences, while worst step refers to the rotation angle which gave the largest number of occurrences. Time Best Step Number of Worst Step Number of Ma degrees Occurrences degrees Occurrences 69.4 47 116 3 135 88.0 64 119 45 137 94.0 37 118 51 136 118.0 28 78 0 93 130.2 37 56 26 69 152.0 17 65 32 80 166.0 59 59 70 70 195.0 43 32 5 42 216.0 24 45 61 56 237.0 45 34 11 42 TABLE 3.7. Comparison of reptile fossil occurrences and predicted upwelling. Shelf area was averaged over the Mesozoic for calculating statistics in Ihe cases of all data lumped together. Reptile occurrences Occurrences Total occurrences Occurrences Total ft reptile Proportion of Age-restrictcd explainable by possibly explainable possibly explainable not explainable occurrences shelf covered Stage upwellingt by upwellingt by upwelling* by upwelling on the shelf by upwelling Statistics Maa 10 10 20 23 43 0.21 0.00010 + Con 1 1 2 1 3 0.23 0.12556 Cen 1 12 13 8 21 0.24 0.00022 + Apt 2 1 3 5 8 0.20 0.14333 Val 0 0 0 4 4 0.26 0.29487 Kim 2 4 6 II 17 0.25 0.12916 Cal 2 2 4 6 10 0.20 0.08731 Pli 0 0 0 3 3 0.28 0.37138 Nor 1 2 3 6 9 0.27 0.25007 Ani n/a n/a n/u n/a n/a Total 19 32 51 67 118 0.24 4.7143643 +

Age-unrestricled Maa 10 10 20 23 43 0.21 0.00010 + Con 12 21 33 30 63 0.23 0.00000 + Cen 5 17 22 15 37 0.24 0.00000 + Apt 5 4 9 14 23 0.20 0.01707 + Val 1 1 2 7 9 0.26 0.29404 Kim 2 7 9 20 29 0.25 0.12261 Cal 5 4 9 10 19 0.20 0.00496 + Pli 1 6 7 15 22 0.28 0.16752 Nor 4 12 16 12 28 0.27 0.00055 + Ani 0 5 5 17 22 0.29 0.16173 Total 45 87 132 163 295 0.24 6.84383 +

t the gnd cell centroid fell within the predicted upwelling area t Ihe grid cell centroid did not fall within predicted upwelling, but some other part of the grid cell did * includes reptile occurrences explainable and possibly explainable by upwelling + the number of reptile occurrences falling within predicted upwelling areas is statistically signitlcanlly different (a=0.05) from the number t^ predicted by the null hypothesis (binomial probabilities calculated for individual stages, G-statislic (G-lest) calculated for all data together) TABLE 3.8. Counts of giid cells containing marine reptile families and their intersection with predicted upwelling. For each data type, the total number of cells containing the specific family is shown first, followed by a column labeled "+UZ", which shows the number of grid cells containing the specific marine reptile family that intersected predicted upwelling. "Cells that intersect upwelling" includes cells containing either lithologic or reptile data; MR = total number of marine reptile occurrences for that time period. Letter next to age indicates age resolution of reptile data; r = agc-restricted; u = age-unrestricted. Family abbreviations arc below the table. Cells diat Age intersect Ma upwelling MR i-ind +UZ i-ich +UZ 69.4-r 49 43 16 9 88-r 54 3 88-u 54 63 20 15 94-r 58 21 6 4 5 2 5 4 94-u 58 37 12 7 13 5 II 8 118-r 24 8 2 0 2 1 3 0 118-u 24 23 8 1 2 1 7 1 130.2-r 15 4 3 0 130.2-u 15 9 4 0 152.2-r 23 17 10 3 7 1 6 2 152.2-u 23 29 15 6 8 1 1 0 13 5 166-r 21 10 3 1 1 1 5 3 166-u 21 19 8 3 6 3 II 86 195-r 10 3 1 0 1 0 10 10 10 195-u 10 22 10 4 6 1 5 1 3 14 1 II 2 216-r 22 9 7 2 1 1 1 0 4 2 216-u 22 28 19 8 I 1 I I 2 1 11 8 6 4 1 0 237-u 12 22 13 3 10 3 9 2 2 0 4 1

Family abbreviations: i-ind = Ichthyosauria indet. or incertae sedis i-ste = Stenopteiygidae i-ich = Ichthyosauridae i-tem = Temnodontosauridae i-lep= Leptopterygidae i-pla = Platypteiygidae i-mix = Mixosauridae p-ind = Plesiosauria indet. or incertae sedis i-sha = Shastasauridae or Shonisauridae p-pto = Pistosauridae TABLE 3,8. (continued)

Age P- P- P- P- P- in- 111- in- m- Marine reptile order totals Ma cry +UZ ela +UZ ple +UZ pli +UZ pol +UZ ind +UZ mos +UZ pli +UZ tyl +UZ ICH +UZ PLE +UZ MOS +UZ 69.4-r IS II 2 1 4 4 19 9 23 9 14 6 3 2 27 16 33 13 88-r 2 0 1 0 1 1 1 1 2 0 2 2 88-u 1 1 26 IS 1 1 14 8 11 6 28 IS 24 11 2S 14 14 9 40 22 51 27 94-r 4 2 3 3 4 4 10 5 13 10 94-u 11 7 8 6 7 6 23 11 24 17 118-r 4 1 2 1 3 1 2 U 4 1 7 2 n8-u 8 4 3 1 5 3 4 2 10 2 18 8 130.2-r 1 0 130.2-u 2 0 2 0 4 1 5 1 5 1 152.2-r 2 0 1 1 7 1 1 0 13 3 12 4 i52.2-u 3 0 1 0 2 1 14 2 1 0 18 6 22 6 166-r 3 1 4 2 4 3 4 2 8 4 166-u 4 2 4 2 2 1 6 4 12 6 12 7 195-r 1 0 1 0 2 0 1 0 195-u S 1 9 3 6 2 15 4 16 5 216-r 1 0 9 3 1 0 216-u 1 0 1 0 25 13 7 4 237-u 20 5 5 1

Family abbreviations: p-ciy = Cryptoclididac m-ind = Mosasaundae indet. or mcertae sedis PLE = Plesiosauna p-ela = Elasmosauridae m-mos = Mosasaurinae MOS = Mosasauridac p-ple = Plesiosauridac m-pli = Plioplatecarpinac p-pli = Pliosauridae in-tyi = Tylosaurinae p-pol = Polycotylidae ICH = Ichlhyosauria K> ON TABLE 3.9 Comparison of reptile fossil occcurrences and predicted upwelling for marine reptile families and orders. Shelf area was averaged over the duration of a taxon for calculating statistics. A) age-restricted data. B) age-unrestricted data. A. Occurrences Occurrences not Total H Proportion of explainable by explainable by reptile shelf covered Taxon Duration* upwellingt upwelling occurrences by upwelling Statistics Ichthyosauria Nor-Cen 14 28 42 0.24 0.0550 family indet. Nor-Cen 10 22 32 0.24 0.1019 Ichthyosauridae Nor-Kim 3 7 10 0.25 0.2505 Leptopterygidae Pli 0 1 1 0.28 0.7200 Mixosauridae Nor 0 1 1 0.27 0.7300 Platypterygidae Apt-Cen 3 4 7 0.22 0.1377 Shastasauridae Nor 2 T 4 0.27 0.2331 Stenopteiygidae Pli 0 1 1 0.28 0.7200 Temnodontosauridae n/a Plesiosauria Nor-Maa 36 35 71 0.24 0.0000 + family indet. Pli-Maa 18 18 36 0.23 0.0003 + Pistosauridae n/a Cryptoclididae Cal-Kim 1 4 5 0.23 0.4058 Elasmosauridae Pli-Maa 16 14 30 0.23 0.0003 + Ptesiosauridae Nor-Apt 2 3 5 0.24 0.2569 Pliosauridae Cal-Maa 9 11 20 0.23 0.0162 + Polycotylidae Kim-Maa 8 4 12 0.23 0.0015 + Mosasauridae Con-Maa 15 20 35 0.22 0.0032 + subfamily indet. Con-Maa 10 10 20 0.22 0.0042 + Mosasaurinae Maa 9 14 23 0.21 0.0239 + Plioplatecarpinae Maa 6 8 14 0.21 0.0391 + Tylosaurinae Con-Maa 3 1 4 0.22 0.0335 + *First and last map intervals over which a taxon is present; intervals within this range were averaged for shelf area tSome part of the grid cell intersected predicted upwelling (explainable and possibly explainable categories of Table 3.7) +The number of reptile occurrences falling within predicted upwelling was statistically significant at the /'=0.05 level (binomial probabilities) TABLE 3.9 (continued). B. Occurrences Occurrences not Total # Proportion of explainable by explainable by reptile shelf covered Taxon Duration* upwellingt upwelling occurrences by upwelling Statistics Ichthyosauria Ani-Cen 48 80 128 0.25 0.0005 + family indet. Ani-Cen 32 57 89 0.25 0.0060 + Ichthyosauridae Nor-Kim 6 15 21 0.25 0.1773 Leptopterygidae Nor-PIi 2 4 6 0.28 0.3137 Mixosauridae Ani-Nor 4 8 12 0.28 0.2191 Platypterygidae Kim-Cen 7 11 18 0.24 0.0697 Shastasauridae Ani-Nor 10 10 20 0.28 0.0201 + Stenopterygidae Pli-Cal 2 2 4 0.24 0.1999 Temnodontosauridae Pli 1 3 4 0.28 0.4177 Plesiosauria Ani-Maa 135 41 176 0.24 0.0000 + family indet. Ani-Maa 50 44 94 0.24 0.0000 + Pistosauridae Ani-Nor 1 4 5 0.28 0.3769 Cryptoclididae Cal-Con 3 5 8 0.23 0.1858 Elasmosauridae Nor-Maa 40 33 73 0.24 0.0000 + Plesiosauridae Nor-Con 7 13 20 0.24 0.1031 Pliosauridae Pli-Maa 27 32 59 0.23 0.0001 + Polycotylidae Kim-Maa 18 9 27 0.23 0.0000 + Mosasauridae Con-Maa 40 44 84 0.22 0.0000 + subfamily indet. Con-Maa 24 23 47 0.22 0.0000 + Mosasaurinae Con-Maa 20 27 47 0.22 0.0009 + Plioplatecarpinae Con-Maa 20 19 39 0.22 0.0000 + Tylosaurinae Con-Maa 11 6 17 0.22 0.0002 + +The number of reptile occurrences falling within predicted upwelling was statistically signiflcant at the P=0.05 level (binomial probabilities) TABLE 3.10. Descriptive statistics for shortest distances between reptile fossil and upwelling-related lithology occurrences for a) age-restricted data, and b) age-unrestricted data. Distances were calculated using Equation 3 (see text) and are expressed in km. "lo" and "hi" refer to the minimum and maximum values for shortest distances for a time interval. a) All lithologies Organic-rich rock Biogenic silica Stage n mean lo hi n mean lo hi n mean lo hi Maa 43 532 0 4016 43 2327 500 6008 43 2750 0 5657 Con 4 444 0 807 4 1435 463 3672 4 1632 0 2296 Cen 21 506 0 4634 21 1866 0 9676 21 2137 491 6204 Apl 8 578 0 1547 8 6753 2186 10519 8 2152 796 3074 Val 4 0 0 0 4 6584 4951 8157 4 1805 0 3229 Kim 17 591 0 2189 17 3550 1035 8433 17 2135 0 6135 Cal 10 2154 0 6706 10 3082 498 6706 10 2713 450 6787 Pli 3 338 0 507 3 762 0 1368 3 1864 1368 2220 Nor 9 1265 0 6027 9 4982 462 16858 9 1959 0 6370 Ani n/a b) Maa (see above) Con 63 814 0 4450 63 4500 0 16678 63 2129 0 6123 Cen 37 505 0 4634 37 2502 0 10846 37 1891 472 6204 Apt 23 1245 0 4689 23 5691 889 10519 23 2627 489 5365 Val 9 1067 0 3705 9 5156 2792 8157 9 2535 0 4980 Kim 29 659 0 2374 29 3512 507 8991 29 1956 0 6135 Cal 19 1771 0 6706 19 3213 498 8953 19 2355 0 6787 Pli 22 1935 0 11393 22 4550 0 13337 22 2975 0 11393 Nor 28 1571 0 6027 28 5176 462 16858 28 2931 0 6593 Ani 22 1556 0 5804 22 5595 0 11060 22 3075 507 11581 TABLE 3.10. (continued) a) Phosphorite Glauconite > 1 lithology/grid cell Stage n mean lo hi n mean lo hi n mean lo hi Maa 43 3037 450 11544 43 1296 0 6390 43 1934 0 6669 Con 4 1576 507 4091 4 1877 1367 2676 4 1087 487 1678 Cen 21 2902 826 6771 21 941 0 5936 21 1014 0 6111 Apt n/a 8 979 482 1678 8 1765 0 3341 Val 4 357 0 955 4 782 482 955 4 1244 0 2271 Kim 17 4762 0 12918 17 1918 0 4804 17 1714 0 6144 Cal 10 7503 2864 14414 10 2991 0 8187 10 2869 0 8022 Pli 3 631 507 879 3 965 507 1510 3 1601 855 2181 Nor 9 7737 2680 17631 9 6330 1825 17824 9 6266 1810 18683 Ani b) Mao (see above) Con 63 2864 0 10698 63 1769 0 7039 63 2178 0 8924 Cen 37 3068 0 6771 37 948 0 5936 37 1175 0 6111 Apt n/a 23 1756 0 5181 23 2391 0 7508 Val 9 4530 0 12299 9 1691 0 4407 9 3226 0 7303 Kim 29 5049 0 12918 29 2173 0 7620 29 1982 0 7278 Cal 19 6796 1229 14414 19 2649 0 8187 19 3090 0 8022 Pli 22 3457 0 13066 22 4084 507 12343 22 4220 0 13249 Nor 28 6761 1379 17765 28 5372 0 17824 28 5430 0 18683 Ani 22 5446 0 15230 22 3832 0 8066 22 .5931 498 14743

K) KJ o TABLE 3.11. Descriptive statistics for shortest distances between reptile fossil and upwelling-related lithology occurrences for a) age-restricted data, and b) age-unrestricted data. Distances were calculated using Equation 3 (see text) and are expressed in km. "to" and "hi" refer to the minimum and maximum values for shortest distances for a time interval. a) Ichthyosauria Plesiosauria Ichthyosaurs + Plesiosaurs* Stage n mean lo hi n mean lo hi n mean lo hi Maa 10 782 0 3228 Con 2 657 507 807 Cen 8 628 0 3696 11 509 0 4634 2 0 0 0 Apt I 0 0 0 4 354 0 895 3 1068 796 1547 Vol 3 0 0 0 1 0 0 0 n/a Kim 5 630 0 1780 4 493 0 1472 8 616 0 2189 Ca! 2 3363 827 5900 6 1633 0 6706 2 2504 0 5009 Pli 2 253 0 507 1 507 507 507 n/a Nor 8 669 0 2680 n/a 1 6027 6027 6027 Ani n/a b) Maa (see above) Con 12 1385 0 4107 Cen 13 725 0 3696 14 469 0 4634 10 268 0 1807 Apt 5 2691 1791 4689 13 800 0 3596 5 959 0 1593 Val 4 1804 0 3705 4 598 0 2391 1 0 0 0 Kim 7 568 0 1780 11 727 0 2374 11 650 0 2217 Cal 7 1935 0 5900 7 1021 0 3137 5 2590 0 6706 Pli 6 1364 0 5594 7 3184 0 11393 9 1343 0 5560 Nor 21 1106 0 3960 3 2377 2046 2663 4 3409 1577 6027 Ani 17 1710 0 5084 2 1690 1522 1858 3 597 0 1790 *Grid cells containing more than one reptile order. These cells were not included in the counts of individual orders. TABLE 3.11. (continued) a) Mosasauridae Plesiosaurs + Mosasaurs* Stage n mean lo hi n mean lo hi Maa 16 656 0 4016 Tt 269 0 Con 2 232 0 463 Cen Apt Val Kim Cal Pli Nor Ani b) Maa Con 22 498 0 3241 29 818 0 4450 Cen Apt Val Kim Cal Pli Nor Ani

to to tv) TABLE 3.12. Number of grid cells containing upwelling-related lithologies and reptile fossils, a) age-restricted data, b) age-unrestricted data. Compare with "c"-group maps in Appendix 1. The next-to-last column contains counts of cells with reptiles and upwelling-related lithologies either coinciding with or adjacent to one another (adjacent == sharing one grid cell side, <=550 km apart). For age-restricted and unrestricted data, a majority of reptile fossil occurrences were associated with upwelling-related lithologies for the Cenomanian and Valanginian map intervals. If reptile-lithology associations one grid cell apart are counted (grid cells share one side, case A of Figure J.I), all map intervals except for the Callovian (restricted and unrestricted), Aptian (unres.), Norian (unres.), and Anisian (unres.) have a majority of reptiles <=550 km away (see Table J. I). a) Coinciding or adjacent # Lithology Marine reptile fossils Upwelling-related lithologies reptile/lithology occurrences Stage + reptile Total # lith+reot Total # lith+reot (distances <==550 km) occurrences occurrences total rept occurrences total lith U cells #cells/total rept Maa 17 43 0.40 92 0.18 32 0.74 Con 1 4 0.25 81 0.01 3 0.75 Cen 15 21 0.71 105 0.14 18 0.86 Apt 3 8 0.38 61 0.05 4 0.50 Val 4 4 1.00 53 0.08 4 1.00 Kim 8 17 0.47 50 0.16 12 0.71 Cal 3 10 0.30 46 0.07 4 0.40 Pli 1 3 0.33 16 0.06 3 1.00 Nor 3 9 0.33 25 0.12 5 0.56 Ani n/a b) Maa (see above) Con 25 63 0.40 81 0.31 37 0.59 Ccn 24 37 0.65 105 0.23 30 0.81 Apt 6 23 0.26 61 0.10 9 0.39 Val 6 9 0.67 53 0.11 6 0.67 Kim 14 29 0.48 50 0.28 19 0.66 Cal 6 19 0.32 46 0.13 8 0.42 Pli 7 22 0.32 16 0.44 12 0.55 Nor 8 28 0.29 25 0.32 II 0.39 Ani 3 22 0.14 15 0.20 3 0.14 TABLE 3.13. Breakdown by lithology type and by reptile order of grid cells containing both reptile fossils and upwelling-related lithologies. a) age-restricted data, b) age-unrestricted data. 0=organic-rich rock, O=biogenic silica, V=phosphorite, Z=glauconite, >1=>I lithology type/grid cell, l=lchthyosauria, P=Plesiosauria, M=Mosasauridae. For each category, cell counts are shown first, followed by the proportion of the total number of lithology + reptile occurrences constituted by that category. a) # Lith + Stage Rept Upwelling-related lithologies Reptile orders cells 0 Q V Z >1 1 P I+P* M P+M* Moa 19 0 0.00 1 0.05 0 0.00 9 0.47 7 0.37 2 0.11 6 0.32 11 0.58 Con 1 0 0.00 1 1.00 0 0.00 0 0.00 0 0.00 0 0.00 1 1.00 0 0.00 Ccn 15 1 0.07 0 0.00 0 0.00 5 0.33 9 0.60 5 0.33 8 0.53 2 0.13 Apt 3 0 0.00 0 0.00 n/a 0 0.00 3 1.00 1 0.33 2 0.67 0 0.00 Val 4 0 0.00 1 0.25 2 0.50 0 0.00 1 0.25 3 0.75 1 0.25 0 0.00 Kim 8 0 0.00 2 0.25 1 0.13 2 0.25 3 0.38 2 0.25 2 0.25 4 0.50 Cal 3 0 0.00 0 0.00 0 0.00 1 0.33 2 0.67 0 0.00 2 0.67 1 0.33 Pli 1 1 1.00 0 0.00 0 0.00 0 0.00 0 0.00 1 1.00 0 0.00 0 0.00 Nor 3 0 0.00 3 1.00 0 0.00 0 0.00 0 0.00 3 1.00 0 0.00 0 0.00 Ani n/a b) Maa (see above) Con 25 6 0.24 2 0.08 2 0.08 6 0.24 9 0.36 2 0.08 0 0.00 12 0.48 11 0.44 Cen 24 3 0.13 0 0.00 2 0.08 8 0.33 11 0.46 7 0.29 9 0.38 8 0.33 Apt 6 0 0.00 0 0.00 n/a 2 0.33 4 0.67 0 0.00 5 0.83 1 0.17 Val 6 0 0.00 2 0.33 2 0.33 1 0.17 1 0.17 2 0.33 3 0.50 1 0.17 Kim 14 0 0.00 5 0.36 1 0.07 4 0.29 4 0.29 3 0.21 5 0.36 6 0.43 Cal 6 0 0.00 2 0.33 0 0.00 2 0.33 2 0.33 2 0.33 3 0.50 1 0.17 Pli 6 2 0.33 1 0.17 3 0.50 0 0.00 1 0.17 1 0.17 1 0.17 4 0.67 Nor 8 0 0.00 4 0.50 0 0.00 1 0.13 3 0.38 8 1.00 0 0.00 0 0.00 Ani 3 I 0.33 0 0.00 1 0.33 I 0.33 0 0.00 1 0.33 0 0.00 2 0.67 *Grid cells containing more than one reptile order. These cells were not included in the counts of individual orders.

lo 4:^ 225

TABLE 3.14.a). Taphonomic states recorded for taxon-localities. The database contains many records of taxa with rather poor preservation and is not biased towards only well- preserved specimens. Database codes used in these queries are shown in parentheses. See also Tables D.4 and G.2. b). Comparison of occurrences containing well-preserved specimens that partially or wholly intersected predicted upwelling and those that did not intersect predicted upwelling. Well-preserved specimens occur both in and out of predicted upwelling areas, so a preservational bias in favor of upwelling areas is unlikely. Numbers shown are sums of occurrences for all stages together. Res. = age-restricted data, Un. = age-unrestricted data.

A. Taphonomic State Number of Taxon-Localities Preservation Articulated individuals (6, 7) 347 Isolated, worn bones (1,2, 3) 96 Not reported (0) 742 Completeness Complete or nearly complete 150 skeleton (a, q) Isolated, unidentified bones or 614 single elements such as teeth (z, e, d, or i) Not reported ($) 175

B. Nearly complete skeleton (a, q) Articulated individuals (6, 7) Data # occurrences # occurrences # occurrences # occurrences set that intersected that fell outside that intersected that fell outside upwelling of upwelling upwelling of upwelling Res. 4 9 12 11 Un. 6 16 44 45 TABLE 3.15. Counts of grid cells containing upwelling-related lithologies and their intersection with predicted upwelling. Total it of grid cells is the number of cells containing lithologic and/or reptile data. For each data type, the total number of cells containing the specific lithology is shown first, followed by a column labeled "+UZ", which shows the number of grid cells containing the specific lithology that intersected predicted upwelling. UL = cell contained any of the 4 kinds of upwelling-related lithologies, 0 = chert (biogenic silica), V = phosphorite, Z = glauconite, O = organic-rich rock. The last 2 columns list the number of grid cells that intersected predicted upwelling and contained both reptile and lithologic data. Cells that Res. Unres. Time Total # intcrscct UL 0 + V + Z + 0 + MR + UL MR + UL Ma grid cells upwelling UL UZ 0 UZ V UZ Z UZ O UZ + UZ + UZ 69.4 116 49 92 41 40 12 19 11 46 24 13 10 12 12 88.0 119 54 81 37 19 6 9 3 22 10 12 10 1 16 94.0 118 58 105 54 36 15 20 10 57 25 23 21 11 18 118.0 78 24 61 21 24 8 4 4 34 13 10 5 3 6 130,2 56 15 53 14 22 6 9 1 25 5 5 3 0 1 152.2 65 23 50 17 28 12 9 0 19 4 5 3 3 3 166.0 59 21 46 16 15 4 7 2 24 8 7 5 2 4 195.0 32 10 16 6 7 3 4 2 5 3 2 0 0 3 216.0 45 22 25 10 20 8 3 1 3 1 3 3 1 4 237.0 34 12 15 9 3 1 5 3 6 3 3 3 n/a 2 227

APPENDIX A MEASURED STRATIGRAPHIC SECTIONS

This appendix contains detailed stratigraphic section data. I measured 5 stratigraphic sections in West Union Canyon, Shoshone Mountains, using a Brunton compass and a Jacob staff (Figure 2.4). In the field, I described each exposed unit and trenched covered intervals. All of the shale units shown on the following graphic columns were trenched. Trenching usually revealed only minor amounts of very weathered shale, and in some cases weathering extended beyond a trenchable depth

(about 0.5 meter).

The sections show observed lithologic and biostratigraphic information and inferred facies assignments (see Chapter 2). Previous work in West Union Canyon includes sections measured by Silberiing (1959) and Hogler (1992a). Permission to work in the area is required from Berlin-Ichthyosaur State Park (Nevada Parks and Recreation) and from the U.S. Forest Service. The Campground section is located on the Grantsville

7.5' Quadrangle (1984), and all other sections are located on the lone 7.5' Quadrangle

(1984). 228

Key to Graphic Columns

Covered Ammonoids interval

Calcareous Bivalves shale uxr\j Lopha cordillerana Wavy McRoberts bedding Brachiopods Lamination O Echinoderms Bivalve laminations Crinoid columnals Fossil hash Gastropods

Trace fossils ^—2 Ichthyosaur bone Burrow mottling

Vertical scale 1 cm=2 meters

Lithologic notes abbreviations Lithology abbreviations lt=light sl=slightly Wacke=wackestone dk=dark Pack=packstone Grain=grainstone Rock colors: weathered/fresh Some colors reported in Munsell notation Strikes reported in azimuth notation

Biostratigraphy abbreviations See Chapter 2 results section l=lower for facies descriptions. m=middle u=upper Notes on graphic columns are C=Camian field observations, not detailed N=Norian petrographic observations. CM=identified by Chris McRoberts DM-J=identified by Danielle Montague-Judd 229

Campground Section

Starting point: From the first hairpin loop in the dirt road leading to the campground at

Berlin-Ichthyosaur State Park, walk perpendicular to strike (034) for a few meters until

limestone float and ledges appear. Section starts above large, fractured argillite ledges of

the clastic member.

Coverage: S facies, includes abundant ichthyosaur occurrences (fossil zones "a" and

"b"ofSilberling, 1959) 230

40 0^0'-O y-9. 0<9 60494-13 Palaeocardiala sp. abundant . -c « •- Argillite, It gray/dk gray

. -• q.'. " 60494-12 Tan/gray

Shale and shell-packed nodules

Sllty, It tan/gray, whole Tossils mostly att^e Argillite, gray/brown-gray, small whole bivalves Discondnuous ledge. It tan/gray 30 «

Shale and shell-packed nodules, 960716-na brown-green/dk gray-brown, scattered shell impressions in shale

/r"r '• .• • Discontinuous ledges; It gray-blue/ ' g •-tt—. •-•• . V-

60494-10 Shale and shell-packed nodules. It gray/dk green-brown Sla Discontinuous ledges; It brown-gray/ Q jw 72995-9.5 dk gray-brown 20 - 60494-9 Argillite, It & dk gray/dk brown-green • Q'. a 60494-8 Especially prominent ledge; It brown- gray/dk gray 60494-7 Argillite, It, dk gray/dk brown-green

60494-6 Lt brown-gray/dk gray

60494-5 Shale and shelly nodules, gray/ gray-brown

60494-4 Lt brown-gray/dk gray, bivalve, 72795-4t echinoder^ and brachiopod hash 72795-4b 1^1- , - -Vl.- 1 72795-3.5 Shale grades into a small pack- stone ledge; shale is It gray/dk gray to 60494-3 Silty, It brown/dk gray

60494-2 Silty, It brown/dk gray Ostreoid bivalves common in first 25 meters of section

Argillite, dk-brown, contains nodules of shell-packed limestone 60494-1 Strike 034 Dip 36 Utkologic notes Biostratigraphy Facies Figure A.1 231

80 Sbl 60494-20 Lt tan-blue/dk gray, hackly weathering, veitlcu joints

60494-19 Silty, gray-tan/dk gray, marly texture

70

60.

Klamathites schucherti muC (DM-J) Sla

50 60494-18 Silty, nodular. It tan/brown-gray, slope forming, discontinuous I^ges, Palaeocardiata sp. abundant

60494-17 Marly, It tan/brown-gray

Marly, It tan-gray/dk gray

60494-15 U tan/dk gray, silty

V -r'j o •yVa'rf o ''>"1 60494-16 Marly, It tan-orange/gray 60494-14 Lt gray/dk gray-brovm, limestone float Biostratigiapby Facies Lithologic notes Figure A.1, continued 232

120 Sbl 71595-27 Blue-gray/dk gray w -v- '• Vif ' •

. 7 71595-26 Pinkish-purple/dk gray; thin, black Scs ?shells

- - • - • 'w) 71595-251 Blue-gray/dk gray, fractured ledges, calcite veins ^undant Sbl ] V 71595-25b Strike 040 dip 31

110 -

Limestone talus

100 —

I V 71595-241 Blue-gray, rust patches/dk gray, Sbl ftactured ledges 1 ^ 71595-24b

90 — 71595-23 Tan/black, hackly weathering, heavily rectystallized

Sbl 71595-22 Tan/black, hackly weathering

Silty, tan/black, slope-former 71595-21

Strike 050 dip 35

Biostiatigiapby Facies Lhtaologic ootes Figure A.1, continued 233

End of section; covered with alluvium; small streambed is 2.5 meters up the section line

Kbl= 140 :D v? 71595-36 Flaggy, tan/dk gray, very small ledge

71595-35 Very weathered brown shale, large & small bone fragments in S=3 72795-35 float Tropites sp. 3 ^^®S=3 uuC (DM-J) Sbl n •viftEna 71595-34 Blue-gray, rust/dk gray, fractured, bone fragments in rock and in float, patches of single shells & hash

Scs 71595-33 Weathered, gray calcareous shale

w E=3 71595-321 Discontinuous outcrops, gray-blue, rust/dk gray, fractured, b^e in float Sbl • 71595-32m and in rock 130 - • •;> fcii! 71595-32b iScsI 71595-31 Very weathered brown shale, possibly calcareous, bone in float :sbi: 71595-30 Blue-gray, rust/dk gray, fractured, slightly fetid, marly iScsI 71595-29 Very weathered brown-black shale, possibly calcareous

71595-28 Blue-gray/gray, fractured, blue-orange mottles Sbl

Biostradgrapby Facies LitboMgic notes Figure A.1, continued 234

Fossil Hill Section

Starting point: On the lone quadrangle, find the side canyon on the north side of West

Union Canyon that begins 3 cm above the top of the "U" in "Berlin-Ichthyosaur".

Section begins between the 7240 and 7280 contours from the base of the west-most stream, which is the top of the clastic member.

Coverage: S facies, includes ichthyosaur occurrences (fossil zones "a" and "b" of

Silberling, 1959) 235

40 70895-3.93 70895-3.92 70895-3.9 70895-3.91 Marly, It tan-giay/dk gray, many whole fossils (especi^ly Palaeocar- diata sp.) in t^us and in rocks; whole fossils most abundant in ledges in the middle of this succession; discon­ tinuous ledges; interfaedded shales 70895-3.8 are noncalcareous and contain rare whole fossils

70895-3.75 Argillite, noncalcareous, gray-brown/ 30 — brown-^een, rare Palaeocardiata sp. impressions, shelly, fetid, limestone 70895-3.5 nodules at base

Klamalhiies sckucherii ^60594-3 Marly, It t^ gra^/dk gray, slope- muC (DM-J) forming, disconanuous ledges, many whole fossils in talus

Argillite, It gray/dk gray 60594-2

s Silty, grainy, tan-gray/dk gray, Sla 60594-1 ledge, gray-tan mottling, some 20 - '•"oQ fossil hash concentrated in gray patches

10

Strike 018 dip 32 S

Biostratigraphy Facies Lithologic nolcs Figure A.2 236

80 60594-9 Lt tan-gray/brown-gray

60594-U.25| Gray-orange mottles Q ^ . 60594-11.1 Blue-black/dk brown, flat laminations

60594-8 Grainy weathering, rust-colored Sbl patches (dolomitic?) near top of bed, possible sponge or coral fiagments 60594-7 Dk blue-brown

70 - CJ? 60594-6 Steel gray/dk steel gray, rust-colored patches, shaly near top, very fi:ac- tured, possible sponge or coral frag­ ments, slope below covered with talus of this lithology

60594-4 Brown, dk brown, blocky

70895-5.2 Marly, whole fossils in talus 60 - ,: A. • q 70895-5.1 Dk gray-brown, slope, splintery Sbl StriM 020 dip 35 S

60594-5 Tan/gray, hackly weathering

:Sla=

SO Whole fossils in talus

iSlail lHy. 70895-3.94 Marly, hash increases from unit 3.8 Biostratigiaphy Facies Lithologic notes Figure A.2, continued 237

120

110 —

70895-13 Gray/dk ^y, rust-colored patches, 60594-13 calcite veins Sbl

100 —

Steel gray/dk gray, set of 3 discon­ tj ? 60594-12a tinuous ledges, Palaeocardiata sp. especially abundant in middle ledge, some whole bivalves and sponge- like (or coralline) fossils, rust- •V -.v.' ] Q? colored splotches 60594-12b

55? 90 — Sbl • 60594-11.5 Lt gray/dk gray, mottled 60594-11 Gray/dk brown, rust-colored patches Steel gray/dark gray, set of 4 beds (packstone at top panly covered), possible coralline fossils

7

60594-10

Biostratigrapby Facies Lithologic note* C0 Figure A.2, continued 238

160

^5? 60594-19L3 Section crosses a right-lateral strike- slip fault; units 19 and 20 are repeats of 13

60594-19L2

60594-20

9 60594-191.1

End of non-repeated section

60594-18 Lt tan-blue/gray, weathers into Scs 140 — stubbly blocks

70895-17.5D Lt gray-tan/gray-brown, bone frag­ ments in float, ammonoids and sinall -1^ 60594-17 :sbi: bivalves especially abundant S=2 60S94-16 Lt brown/dk gray-black Scs 70895-15.5 Lt brown/dk gray-black Lt gray-tan/gray-brown, bone frag­ :sbi: 70895-17.50 ments in float, ammonoids and si^l bivalves especially abundant 130 —

60594-15 Lt brown/dk gray-black, breaks into 70895-15 shards Lt gray-tan/gray-brown, bone frag­ :sbi: ft pa q- o' .'.'S Q 1 70895-17.n ments in float, ammonoids and small bivalves especially abundant 60594-14.5 Scs 60594-14 G^ay^rown-g^ay

120 Biostratigiapby Facies Lithologic notei Figure A.2, continued 239

Boundary Hill Section

Starting point: On the lone quadrangle, find the two small ridges that intersect the northern boundary at the northeast comer of Berlin-Ichthyosaur State Park. Section starts at the base of the northem ridge, directly along the park boundary. Ammonoids (Tropiies sp.) and ichthyosaur bones are abundant over the first 20 meters of section. Outcrops are thin and sparse over the remainder of the section.

Coverage: Sbl, Scs, and Pel subfacies, includes ichthyosaur occurrences (fossil zones

"b"'and "c" of Silberling, 1959) 240

40

5=3

E=a

30 - 5=3 71095-4 Float-limestone talus %

] 71095-3 Gray, orange/dk gray, many Tivpites sp. Tropites sp. 3S=3 Q uuC (DM-J) Sbl ] (S 71095-2 Lt blue-lan-orange/dk gray, many 9607-1 bivalves (external mol&), Tmpites sp.; bone in float

20 -

Especially dense concentration of bone Gragments in limestone talus; 71095-1 f many Dvpites sp. and small bivalves

5=3

10 - 5=3

Tropites sp. S=3 uuC (DM-J) %

Strike 041, dip 20

Biostratigiaphy Fades LHhologic notes Figure A3 241

80

!Scs 71095-11 Lt gray, puiple, giainy weathering

71095-10t Gray/dk gray, discontinuous ledges, Sbl=| 73095-lOt trenching yielded dirt, calcite, and limstone talus, reddish alteration patches, grainy weathering ^Sbl: 73095-10m

70 71095-10b Strike 065, dip 30 Sbl=1 73095-1 Ob

60 -

Sbl Silty, tan-red-pink, fiactured

=Scs= 71095-6 Gilcareous, tan-red S=3 71095-5 Tan-red/dk gray, intensely fractured -Sbl and possibly hy^othennally altered (red patches)

40 Biostiatigiaphy Facies Lithoioglc notes Figure A J, continued 242

120

Biown-^y-green/blaclc, grainy weathering, discontinuous, fractured rSbl= 71195-24 ledge . 73095-23 71195-23 Purplish-red, blocky 71195-22.5

T- T T T -r -r -rS 71195-22 Brown-gray-purple/black, discontin­ •(§,? uous, si fetid, grainy weathering Scs 71195-21 71195-20 Gray-purple/black, discontinuous, fractured grainy weather)ng,sl fetid 71195-19 Red-pink, discontinuous 110 - 71195-18 Lt gray-purple/blacl^ discontinuous, fiacturcd ledge, grainy weathering, si fetid, tiny ammonoids, rare bivalve impressions rScsz 71195-17 Grainy weathering

100 — :Sbl= 71195-16t Gray-brown/dk gray, grainy weather­ :Sbl= ing, si fetid :Scs:: 71195-15.5 Red, pink

71195-151 Brown-gray-green/dk gray, set of =Sbl= 3 snull ledges, grainy weathering, si fetid

=SbI= 7U95-15m

=Sbl= 71195-15b 90 73095-15 :Scsi 71095-14.5 Reddish, soily

:Sbt 71095-14 Gray-brown/dk gray, si fetid, fractured iScsI 71095-13 Pinkish brown, si fetid :Sbl= 71095-12t Brown-^y/dk gray-black, grainy weathering, si fetid 71095-12b Stnke027,dip26 ISbl: 73095-12

80 Biostraugiaphy Facies UtholoKic notes Figure A J, continued 243

160

=Scs- 71195-29 Pinlc-brown, weathered, si fedd

:Scs= 71195-28 Gray-brown, discontinoiu, tactured, si fetid 71195-27 Calcareous shale talus

150 -

71195-26 Blue-gray/black Is talus, probably float blocks &om rocks above the section

140 —

71195-25 Blue/dk gray, white recrystallized Pel?: shells 130 —

120 Biostiatigiaphy Facies LHlialogic oolcs Figure AJ, continued 244

200

Guembelites sp. UN (DM-J) Sbl

Guembelites sp. 71195-31 Giay/black, discontinuous ledges, IIN (DM-J) Sbl many Guembelites sp., hashy Halobia sp., gnuny weathering, ammonoi^ easiest to find in float 190 -

Guembelites sp. UN (DM-J) Sbl

Guembelites sp. IIN (DM-J) Sbl 180 -

160 Biostratigiaphy Fades Litbologic Doles Figure A3, continued 245

240 (36 meters to beginning of Brachio- pod Ridge section; all 36 meters covered)

End of measured rock units; section extended to beginning of Brachiopod Ridge section £ong strike 029, dip 29. 210 —

71195-32 Gray-blue/dk gray-black, fetid, Sbl calcite veins

Cray/black, discontinuous ledges, many Guembelites sp., bashy Guembelites sp. 71195-31 Halobia ^., grwy weathering, IIN (DM-J) Sbl ammonoi^ easiest to find in float

Biostratigraphy Facies Lithologic note* Figure A J, continued 246

Furnace Hill Section

Starting point: On the lone quadrangle, draw a line from the symbol marking the

northeast comer of Berlin-Ichthyosaur State Park north to the 7360 contour on the north

side of Union Canyon. Section starts about 5 meters up from the gully that is at the base

of the west-facing slope.

Coverage: Scs and Pel subfacies (fossil zone "c" of Silberling, 1959) 247

40 60694-17 Lt brown, flaky Scs 73095-17 Section interrupted by felsic dikes

^ 60694-15 Dk blue-raay/dk gray, thin-medium Pel L. 72995-15 bedded, fossil hash laminations at top

-Sbl- 60694-14 Pinkish, irregular laminations in cal­ .Scs. 73095-14 careous shale 30 - 60694-13 Lt blue-gray/dk gray, laminations

:Sbl= 60694-12 Platy :Scs 60694-11 Brown

iScsI 60694-10 Lt brown-gray/brown, splintery 9> 9 9 60694-8 Lt brown/brown, platy 20 72995-lOg :SbI: 72995-1 Of

72995-lOe Scs 9 72995-10c,d :Sbb 72995-10b iScs= 72995-lOa 9 iScsi 69694-5 Lt giay/dk gray Halobia cf. 60694-4 Lt blue/dk gray, grainy weathering beyrichi (Mojsis- ovics) Sbl 60694-3 IN (CM) 72995-1.5 to 60694-1 Lt gray/dk gray-black, wavy lamin­ 72995-1 ations Scs

60694-2

Strike 015-020, dip 33 0 I Lithologic aolet BiostratigTaphy Facies 58/ V Figure A.4 248

80

60694-27 72995-27b Sbl 72995-27a Scs 60694-26 Tan-orange/dk brown-blaclc, blocky

60694-25 Steel gray/dk gray, clifT-fonner, silty, 72995-25 lammateo patches, homfelsed Pel 70 -

Scs 72995-24.5 Lt purple shaly soil

60694-24 Gray-tan-purple Scs 72995-24 72095-24

Halobia cf. ISbl 72995-23.5a,b Tan-gray/dk gray beyrichi (Mojsis- 72995-23.2 ovics) :SbN 60 72995-23.1 Lt blue/dk gray IN (CM) sbr Orange-gray/dk gray, bed coarsens Guembelites sp. 60694-23 upward, altered limestone; many UN (DM-J) Sbl 72195-23 ammonoids in float in the interval between samples 21 and 23.5

60694-21 Lt blue-^y/dk blue-gray, altered 72995-21C (felsic dike interrupts section between Sbl 72995-2lb units 21 and 23) 72995-2la 60694-20 Weathers purple-blue, blocky, contains -Scs! 73095-20 limestone nodules

Blue-gray/dk gray, medium bedded, 60694-19 grainy weathering, silty laminations, sharp, undercut contact at base, coarsens upward Pel 50 - 72995-19

60694-18 Blue-gray/dk gray, medium bedded, Halobia cf. Pel 72995-18 grainy weathering, silty laminations, beyrichi (Mojsis- coarsens upward ovics) jf jpjpjpjp jp *j" 60694-18a IN (DM-J) Scs Pet Scs 73095-17J Lt brown, flaky, top contact with limestone is sharp

40 Biostratigraphy Facies Lltbologic notes Figure A.4, continued 249

110 -

60694-29 Blue-dk orange/gray, calcite veins, Pel 72995-29 small-scale fold iScsZ 72995-28.5 Brown/black

Scs

=Sbl= 72995-2i Shales-brown/black Wsckestones-biuc/dk gray Shales get more calcareous up-unit Scs 100 and conlain limestone nodules; limestone ledges discontinuous :Sh\z IScsI :Sbli

Halobia cf. 60694-28b beyrichi (Mojsis- Scs ovics) IN (DM-J)

90 —

80 Biostiatignipby Fades Litliologic aotes Figure A.4, continued 250

160

150 -

End of section-obscured by volcanic outcrops and talus

140 — Blue-gray/dk gray, cliff-former, rare silicified brachiopods, large calcite veins, wavy, nonpwallel bedding (medium thickness)

Pel

130 — 60694-30 72095-30 72995-30

Volcanic talus

Biostratigiaphy Fades Litbolosic notes Figure A.4, continued 251

Brachiopod Ridge Section

Starting point: On the lone quadrangle, find the eastern boundary to Berlin-Ichthyosaur

State Park. Section starts at the 7360 contour just east of the park boundary and along the northwest-facing slope of the large ridge intersects the eastern park boundary (near the

7335T benchmark).

Coverage: Pel, Pbl, Sbl, O, and F subfacies 252

40 60794-2a Dk blue-giay/dk gray, laminated

O 60794-2b Dk blue-gray/dk gray

30 -

Gray/dk blue-gray

O

60794-2 20 —

10 —

N4/N2^, wavy, nonparallel bedding 60794-1 o (medium thiclmess), calcite veins Strike ISS, dip 37

Biostratigiaphy Fades Litbologic notes Figure A.5 253

3>bi: 3»bi:

yb\: 60794-8 N4.S/N3, discontinuous ledges

N4/N3, nonparallel wavy bedding, some silicined patches

73095-7m N5.5/N3.5

Pbl

60794-7

60794-6 NS/N2.S, nonparallel wavy bedding, many calcite veins, localized brown (chert?) nodules

60794-5 N4/N3, nonparallel wavy bedding, thin-mediumbedded, calcite veins, Pbl fetid, base undercut

o 60794-3 N6-N4/N2, massive, calcite veins

Strike 135, dip 40

Q Dk blue-gtay/dk gray, thinly bedded

Biostratigiaphy Facies Lithologk notes Figure A.5, continued 254

120

iScs: 60794-13 SY2/1, fossiliferous olive-black shale

60794-9 N6/N3, cliff-former, dense patches 73095-9 (30-40 cm diameter) of silicified 90 brachiopods and hash

Pbl

PHT top ledge of unit 8—see previous page

80 Biostradgiaphy Facies Lithoiogic aotes Figure A^, continued 255

160

960719-15 Silty, f1ag», abundant whole biach- iopods, ledge, top contact sharp and -(Kii ij^.. 73195-0.5 iiregular __±_1 -tr - •

•fr -ir o 7 . ••«: -A- 60794-15 N5-4/N3, clifT-fonner, thin-medium

iScs: 60794-14 Brown/black, fine laminations

60794-11 N2.S-3/N2, somewhat fetid, under­ Sbl cut base

60794-10 Lt olive-gray. It med bIuish-gray/N3, Scs 73095-10 hashy p<^ets, thin-medium beds 60794-12 Black, laminated

Biostratigraphy Facies LUhologic notes Figure A.5, continued 256

200 jW .y\A» vTvA/ J\/^

sj\n^ 60994-17 N3.5/N2.5, eroded ledge, ostreoid - 73195-17 bivalves common, hackly weathering N4.S/N3, cliff, grainy, top contact 60994-16 irregular, hackly weathering 60994-15.5 X»'- 60994-15 5B 5/1 (weathered)/N3 (fresh), dis­ continuous la^e ledges, nodular, base less fossiliferous than top, fetid Lopha cordillerana 60994-14 McRoberts <1 N (DM-J) ^7—V 0 60994-13 N5/N3, discontinuous large ledges, ^ ^ o O ^ 3 70795-13 spv-fllled brachiopods, some fossils 190 — "TT— ''' ^ silicified _.-'p .V 0 Strike 025, dip 28 f -SSi lV O n .AA. ^ —- , vw -/ r *•/ 601^94-12 N7/N2.5, cliff, calcite veins, hackly weathering

60994-11 N7/N3, cliff, base undercut, nodular, S) hackly weathering

180 — 60994-10 N6/N3, cliff

60994-9 N6-7/N2.5, cliff 60994-8 N6/N3, cliff, grainy 60994-7 N6/N3, undercut, nodular texture 60994-6 N6/N3, cliff, a few whole ribbed shells

60994-5 N6/N3, cliff, massive

60994-4 N6/N3, cliff, massive, hackly weathering

170 — 60994-3 N7(4yN2.5 (top), N3-2.5/N2 (base) 73195-3 cliff, undercut, nodular, hackly weathering

oo ^ o o ^ i' 60994-2 N2/NI.5, undercut ledge, fetid '•'Vv O 73195-2 Strike 360, dip 28 o o o

*' 60994-1 N5/N3.5, cliff, grainy, fetid, grains o partially silicified 160 Biostiatigiaphy Fades LitholoKic notes Figure A.5, continued 257

NS/N3, exposed intermittently, medium-b^ed ledges, patchy fossil exposure, some burrow mottles filled with coarser material

(m float)

N7-NS<'N3, cliff, dip-slope, thickly bedded with a few thin, fetid beds that are silty and undercut, hackly weathering; whole brachiopods erode out of undercut beds, sponge­ like fragments present

Lopha cordillerana 220 — McRoberts 60994-24 N (CM, unit 21; DM-J, other units) NS.5/N2, cliff, massive, hackly weathering

60994-22 N5/N2.S, cliff, massive, hackly weathering

N5.5/N2, cliff, hackly weathering

N6.5/N2.5, cliff, thickly bedded, undercut at top, ThalassonoUies-type trace fossils at base, ostreoid bivdves 60994-21 concentratnl at base and top of bed 73195-19-21 60994-19 Med. olive gtay/N2.S, undercut mmt N2/N2, undercut, fetid (unit 18)

NS-7/N2, cliff, pockets of fossil hash, 60994-17.5 ostreoid bivalves concentrated at top, 73195-17.5 in undercut sediments

Biostratigraphy Fades Lithologic notes Figure A3, continued 258

280 Interbedded shales and limestones, biachiopods in limestoc^ are larger than those lower in the section (unit 24 and below), limestone beds thin upsection, individual beds coarsen upwards and have hashy tops

XI

270 -

260

•«sT 60994-27

250 "5] 60994-26 60994-25

Lopha cordillerana McRobots N (DM-J)

Biostiatigiaphy Fades Lithologic notes Figure A.5, continued 259

320

310 - End of section; beds upsection are covered by volcanic rocks

60994-30 Gray/dk gray, ledges, nuiny brachio- pods in the hsshy intervals 1 '

300 — 60994-29

60994-28

290

Biostiatigiaphy Facies LHlialogic natci r ^ Figure A^, continued 260

APPENDIX B

RESULTS OF INORGANIC AND TOC ANALYSES FOR SAMPLES FROM THE LUNING FORMATION AT WEST UNION CANYON 261

TABLE B. 1. Raw results for TOC analysis. Replicate samples were analyzed in a separate batch and their TOC values were consistently lower than the first batch, perhaps in part because of instrumental drift. For this reason, average TOC values for replicated samples were not calcaluated. "Meters" refers to relative position in the composite stratigraphic section. Key to stratigraphic sections (Appendix A): Camp=Campground, FoH=Fossil Hill, BH=Boundary Hill, Fum=Fumace Hill, BR=Brachiopod Ridge.

Field Number Section Meters Lithofacies TOC, wt % 604941-1 Camp 5 Sla 0.19 708951-3.93 Camp 40 Sla 0.09 708951-3.93 Camp 40 Sla 0.01 Replicate 708951-3.93 Camp 40 Sla 0.02 Replicate 605941-8 FoH 75 Sbl 0.29 604941-20 Camp 80 Sbl 0.23 72795-15 FoH 128 Scs 0.51 72795-15 FoH 128 Scs 0.34 Replicate 605941-17 FoH 135 Sbl 0.10 71095-3 BH 138 Sbl 0.18 73095-10 BH 190 Sbl 0.35 73095-12 BH 198 Sbl 0.34 71095-14 BH 201 Sbl 0.42 73095-15 BH 212 Sbl 0.26 71195-23 BH 228 Scs 0.81 72995-1 Fum 232 Scs 0.85 72995-1 Fum 232 Scs 0.65 Replicate 72995-1 Fum 232 Scs 0.76 Replicate 72995-8-10 Fum 243 Scs 0.46 72995-lObc Fum 248 Sbl 0.38 72995-15 Fum 257 Pel 0.19 71195-28 BH 267 Scs 0.41 72995-18 Fum 268 Pel 0.11 72995-19 Fum 274 Pel 0.21 72995-2 lb Fum 278 Pel 0.28 72995-25 Fum 293 Pel 0.20 72995-27a Fum 297 Scs 0.24 71195-32 BH 319 Pel 0.24 606941-30 BR 359 Pel 0.08 607941-6 BR 423 Pbl 0.01 70795-15 BR 513 O 0.06 73095-0.5 BR 514 O 0.06 73195-2 BR 522 F 0.24 73195-2 BR 522 F 0.12 Replicate 609941-18 BR 559 F 0.11 609941-28 BR 650 F 0.04 262

TABLE B.2. Raw results for ICP analysis. "Meters" refers to relative position in the composite stratigraphic section. Key to stratigraphic sections (Appendix A): Canip=Campground, FoH=FossiI Hill. BH=Boundary Hill. Fum=Fumace Hill. BR=Brachiopod Ridge. Italicized elements might have been incompletely extracted: underlined elements might have been lost during extraction. Replicate samples were averaged (denoted by A (-'G columns) and used in Figures 2.14 and 2.15 and Table 2.5. Field numbers ending in "i" were samples of bivalve internal molds.

Section Camp Camp FoH FoH FoH Camp FoH Field 604941- 604941- 708951- 70895- 70895- Arc 604941- 605941- number 1 12i 3.9i 3.93 3.93 20 8 Element Detection Facies Sla Sla Sla Sla Sla Sla Sbl Sbl limit Meters 5 37i 40i 40 40 40AVG 80 75 Be ppm 0.5 0.5 <0.5 <0.5 <0.5 <0.5 <0.5 <0.5 0.5 Na wt% 0.01 0.17 0.06 0.16 0.08 0.07 0.08 0.06 0.05 Mg wt % 0.01 0.22 0.47 0.41 0.34 0.31 0.33 0.88 0.30 Al wt % 0.01 1.67 1.15 1.34 0.79 0.70 0.75 1.88 0.62 P wt% 0.01 0.08 0.09 0.06 0.05 0.06 0.06 0.03 0.05 Kwt% 0.01 0.31 0.05 0.34 0.14 0.12 013 0.11 0.02 Ca wt% 0.01-30 >30 12.40 >30 >30 >30 >30 >30 >30 Sc ppm 0.5 4.6 5.5 3.3 2.7 2.0 2.4 3.5 1.3 Ti wt % 0.01 0.07 0.02 0.05 0.03 0.03 0.03 0.08 0.03 y ppm 2 25 20 24 14 12 13 27 10 Cr ppm 1 20 23 15 14 13 14 18 18 IVIn ppm 2 1340 1500 1590 900 842 871 516 796 Fe wt % 0.01 1.91 1.77 1.18 1.19 1.17 1.18 1.23 1.50 Co ppm 1 5 8 4 1 2 2 3 1 Ni ppm 1 9 58 18 8 7 8 18 15 Cu ppm 0.5 8.7 79.9 8.7 5.4 4.2 4.8 13.0 9.6 Zn ppm 0.5 45.6 56.6 27.1 28.6 26.6 27.6 56.6 53.0 As ppm 3 8 <3 <3 <3 <3 <3 <3 4 Sr ppm 0.5 336 144 472 631 571 601 633 972 Y ppm 0.1 10.4 11.1 II.7 10.1 10.4 10.3 4.8 2.6 Zr ppm 0.5 12.0 9.9 19.8 9.3 10.0 9.7 23.1 7.1 Mo ppm 1 1 <1 2 2 2 2 9 7 Ag ppm 0.2 <0.2 0.5 <0.2 <0.2 <0.2 <0.2 <0.2 <0.2 Cd ppm 1 <1 <1 <1

TABLE B.2., continued

Section FoH FoH BH BH BH BH BH Field 72795-15 72795-15 AVG 71095-3 71095-11 73095-15 71195-18 73095-23 number Facies Scs Scs Scs Sbl Scs Sbl Scs Scs Meters 128 128 I28AVG 138 196 212 220 228 Be ppm 1.2 1.2 1.2 <0.5 0.7 <0.5 0.5 0.7 Na wt % 0.55 0.54 0.55 0.07 0.17 0.07 0.12 0.17 Mg wt % 0.40 0.30 0.35 1.11 1.08 1.06 0.96 1.09 Al wt % 5.24 5.38 5.31 0.43 2.31 0.76 0.85 2.0! P wt % 0.09 0.10 0.10 0.05 0.06 0.04 0.04 0.12 K wt% 1.00 1.05 1.03 0.08 0.49 0.16 0.22 0.42 Ca wt % 21.20 20.30 20.75 >30 27.10 >30 >30 >30 Sc ppm 10.5 10.2 10.4 0.8 4.6 1.7 2.2 3.8 Ti wt % 0.23 0.24 0.24 0.02 0.08 0.03 0.04 0.08 y ppm 76 76 76 9 111 24 21 133 Cr ppm 57 33 45 14 28 10 13 31 Mn ppm 669 456 563 397 157 204 215 179 Fe wt % 3.78 2.82 3.30 1.05 0.83 0.47 0.68 1.20 Co ppm 23 3 13 <1 1 <1 <1 3 Ni ppm 155 23 89 11 28 12 14 53 Cu ppm 47.6 17.7 32.7 4.9 17.7 5.5 9.2 21.4 Zn ppm 64.3 66.5 65.4 42.1 121.0 73.7 76.7 223.0 As ppm 7 <3 <3 <3 14 3 <3 8 Sr ppm 648 650 649 1230 1510 1270 1140 1860 Y ppm 12.1 12.3 12.2 2.8 13.7 4.9 4.6 12.5 Zr ppm 50.7 54.9 52.8 10.0 24.1 14.4 16.8 23.8 IVIo ppm 4 2 3 2 8 2 2 7 Ag ppm 0.2 <0.2 <0.2 <0.2 <0.2 <0.2 <0.2 <0.2 Cd ppm <1 <1 <1 <1 8 2 <1 6 Sn ppm <10 <10 <10 <10 <10 <10 <10 <10 Sb ppm <5 <5 <5 <5 <5 <5 <5 <5 Bappm 251 262 257 23 146 43 62 116 La ppm 15.0 15.4 15.2 2.6 6.4 2.8 3.3 6.1 yy ppm <10 <10 <10 <10 <10 <10 <10 <10 Pb ppm 4 7 6 6 3 3 2 <2 Bi ppm <5 <5 <5 <5 <5 <5 <5 <5 264

TABLE B.2., continued

Section BH BH Fum Fum Fum Fum Field 71195-23 71195-23 AVG 72995-1 72995-1 72995-1 Arc 606941-3 number Facies Scs Scs Scs Scs Scs Scs Scs Sbl IVIeters 228 228 228AVC 232 232 232 232AyG 234 Be ppm 0.6 0.7 n.7 0.7 0.7 0.7 0.7 <0.5 Na \vt % 0.17 0.18 0.17 0.23 0.24 0.24 0.24 0.14 Mg wt% 0.95 0.99 I.OI 0.35 0.35 0.35 0.35 0.44 Al wt% 1.68 2.11 1.93 2.26 2.24 2.14 2.21 1.03 P wt% 0.12 O.iO O.ll 0.12 0.11 0.12 012 0.05 Kwt% 0.36 0.38 0.39 0.39 0.39 0.39 0.39 0.14 Ca wt % 28.60 >30 >30 >30 >30 >30 >30 >30 Sc ppm 3.8 3.9 3.8 4.2 4.2 4.3 4.2 1.7 Ti wt % 0.08 0.08 0.08 0.09 0.09 0.09 0.09 0.04 y ppm 96 99 109 41 43 41 42 25 Cr ppm 28 26 28 29 32 25 29 15 Mn ppm 169 169 172 195 193 195 194 250 Fe wt % 0.90 0.99 1.03 l.ll 1.32 1.10 1.18 0.85 Co ppm 1 I 2 1 5 <1 ri'a <1 Ni ppm 30 32 38 19 52 19 30 15 Cu ppm 20.1 18.2 19.9 17.0 26.2 17.0 20.1 7.8 Zn ppm 250.0 255.0 242.7 66.0 66.4 67.1 66.5 42.7

As ppm 6 6 '/ <3 3 <3 n/a <3 Sr ppm 1670 1820 1783 1750 1710 1750 1737 1470 Y ppm 9.7 9.5 10.6 9.3 9.2 9.3 9.3 4.0 2fr ppm 20.6 22.6 22.3 24.9 20.6 24.7 23.4 19.7 Mo ppm 5 5 6 2 3 2 2 2 Ag ppm <0.2 <0.2 <0.2 <0.2 <0.2 <0.2 <0.2 <0.2 Cd ppm 3 3 4 2 1 2 2 <1 Sn ppm <10 <10 <10 <10 <10 <10 <10 <10 Sb ppm <5 <5 <5 <5 <5 <5 <5 <5 Bappm 99 103 106 108 108 107 108 41 La ppm 5.1 6.0 5.7 8.4 8.0 8.1 8.2 4.0 W ppm <10 <10 <10 <10 <10 <10 <10 <10 Pb ppm 3 3 3 3 4 <2 n/a 3 Bi ppm <5 <5 <5 <5 <5 <5

TABLE B.2., continued

Section Furn Fum Fum Fum Fum BH Fum Field 73095-8-10 73095-lOg 72995-1 Obc 72995-18 72995-18 Ai'G 71195-32 72995-28 number Fades Scs Scs Sbl Pel Pel Pel Pel Scs Meters 243 248 248 268 268 268AyC 319 327 Be ppm 0.8 0.7 0.8 <0.5 <0.5 <0.5 <0.5 0.6 Na wt % 0.19 0.26 0.23 0.04 0.03 0.04 0.09 0.16 Mg wt % 0.36 0.35 0.36 0.34 0.35 0.35 0.39 0.41 Al wt % 1.65 2.62 2.33 0.20 0.21 0.21 0.67 1.25 Pwt% 0.03 0.07 0.05 0.03 0.03 0.03 0.03 0.04 K wt% 0.28 0.55 0,47 0.03 0.03 0.03 0.15 0.33 Ca wt % >30 28.50 >30 >30 >30 >30 >30 >30 Sc ppm 3.4 5.1 5.4 0.5 0.5 0.5 1.5 3.1 Ti wt % 0.06 0.11 0.10 <0.01 <0.01 <0.01 0.03 0.06 yppm 54 136 181 31 33 32 19 32 Cr ppm 29 30 18 8 8 8 10 15 Mn ppm 339 320 341 154 160 157 174 379 Fe wt % 1.81 1.36 0.75 0.30 0.31 0.31 0.47 1.03 Co ppm 1 2 2 <1

TABLE B.2., continued

Section Fum Fum BR! BRI BR2 BR2 Field 606941-30 606941-30 Al'G 607941-6 70795-15 73195-2 73195-2 .•WG number Fades Pel Pel Pel Fbl 0 F F F Meters 359 359 359AVC 423 513 522 522 522AVG Be ppm <0.5 <0.5 <0.5 <0.5 <0.5 <0.5 <0.5 <0.5 Na wt % 0.04 0.05 0.05 0.16 0.04 0.04 0.05 0.05 Mg wt% 0.39 0.32 0.36 0.26 1.03 0.64 0.60 0.62 Al wt % 0.28 0.29 0.29 0.57 0.44 0.80 0.69 0.75 Pwt% 0.02 0.02 0.02 0.04 0.04 0.03 0.03 0.03 K wt % 0.07 0.09 0.08 0.24 0.20 0.36 0.30 0.33 Ca wt % >30 >30 >30 >30 >30 >30 >30 >30 Sc ppm 0.5 0.6 0.6 1.6 1.7 1.5 1.3 1.4 Ti wt % 0.01 0.01 0.01 0.02 0.02 0.03 0.03 0.03 y ppm 8 7 8 12 13 16 15 16 Cr ppm 6 6 6 8 11 9 8 9 Mn ppm 117 93 105 230 357 186 181 184 Fe wt % 0.19 0.20 0.20 0.13 0.96 0.42 0.41 0.42 Co ppm <1 <1 <1 <1 4 <1 2 rUa Ni ppm 9 7 8 8 14 12 16 14 Cu ppm 3.4 2.4 2.9 4.5 4.8 4.6 7.1 5.9 Zn ppm 15.2 36.8 26.0 14.4 9.8 26.9 20.1 23.5 As ppm <3 <3 <3 <3 7 <3 <3 <3 Sr ppm 3290 3250 3270 736 368 773 813 793 Y ppm 1.7 1.4 1.6 14.6 8.3 3.3 3.0 3.2 Zr ppm 10.1 8.2 9.2 16.4 7.1 8.8 7.4 8.1 Mo ppm 2 <1 na 1 3 3 4 4 Ag ppm <0.2 <0.2 <0.2 <0.2 <0.2 <0.2 <0.2 <0.2 Cd ppm <1 <1 <1 <1 <1 <1 <1 <1 Snppm <10 <10 <10 <10 <10 <10 <10 <10 Sb ppm <5 <5 <5 <5 <5 <5 <5 <5 Ba ppm 32 36 34 12 14 13 14 14 La ppm 1.8 1.4 1.6 5.9 5.0 3.0 2.8 2.9 W ppm <10 <10 <10 <10 <10 <10 <10 <10 Pb ppm 4 3 4 7 12 2 4 3 Bi ppm <5 <5 <5 <5 <5 <5 <5 <5 267

TABLE B.2., continued

Section BR2 BR2 Reference Field 609941-18 609941-28 SGR-I SGR-1 AVC number Fades F F n/a n/a n'a IVIeters 559 650 n/a n/a AVC Be ppm <0.5 <0.5 1.1 1.1 I.I Na wt % 0.03 0.03 2.22 2.86 2.54 Mg wt % 0.26 0.27 2.66 2.58 2.62 Al wt % 0.34 0.56 3.15 3.52 3.34 Pwt% <0.01 0.0! 0.12 0.13 0.13 Kwt% 0.15 0.25 1.36 1.40 1.38 Ca wt % >30 >30 5.79 5.77 5.78 Sc ppm 1.5 1.2 4.5 4.4 4.5 Ti wt % <0.01 0.02 0.13 0.14 0.14 y ppm 8 11 123 102 113 Cr ppm 7 16 40 47 44 Mn ppm 129 252 232 239 236 Fe wt % 0.13 0.72 1.96 1.98 1.97 Co ppm <1 8 10 II II Ni ppm 9 58 37 37 37 Cu ppm 3.2 15.9 64.3 64.0 64.2 Zn ppm 19.6 18.9 88.7 100.0 94.4 As ppm <3 <3 17 <3 n/a Sr ppm 463 775 347 348 348 Y ppm 3.8 2.9 9.2 9.4 9.3 Zr ppm 8.5 7.4 37.9 191.0 114.5 Mo ppm 2 3 33 33 33 Ag ppm <0.2 <0.2 <0.2 0.5 n/a Cd ppm <1 <1 <1 <1 <1 Sn ppm <10 <10 <10 <10 <10 Sb ppm <5 <5 <5 <5 <5 Ba ppm 8 9 271 276 274 La ppm 2.3 2.7 18.8 18.4 18.6 iV ppm <10 <10 <10 <10 <10 Pb ppm 3 5 39 35 37 Bi ppm <5 <5 <5 <5 <5 APPENDIX C

DATA USED FOR FIGURE 2.16 269

TABLE C.1. Abundance data for Section 1 of Hogler (1992a). compiled from Appendix 2 of Hogler (1992a). Data are displayed by lithofacies in Tables C.2 and C.3 and in Figure 2,16. Section 1 of Hogler (1992a) spans the marine part of the tuning Formation exposed in West Union Canyon and thus includes the lithofacies described in Chapter 2 of this dissertation. Sample size was not reported in Hogler (1992a). Biofacies-community* and locality and sample numbers from Hogler (1992a).

Taxon Number of Biofacies- Locality Sample Individuals Community Number Number Costatoria sp. 2 l-L 11195 M Halobia sp. 1 1-L 11195 1-1 indet. nautiloid 1 l-L 11195 I-I Lopha montis-caprilis 18 l-L 11195 I-l Septocardia sp. 11 l-L 11195 1-1 snail A 9 l-L 11195 I-I snail W 1 I-L 11195 I-l Zugmayerella sp. 1 l-L 11195 1-1 Halobia sp. 3 l-L 11196 1-2 indet. colonial coral 2 I-L 11196 1-2 Lopha montis-caprilis 41 l-L 11196 1-2 Septocardia sp. 42 l-L 11196 1-2 snail A 12 I-L 11196 1-2 snail W 7 I-L 11196 1-2 decapod 1 1-S 11197 1-3 indet. nautiloid 4 1-S 11197 1-3 Lopha montis-caprilis 23 I-S 11197 1-3 Septocardia sp. 201 I-S 11197 1-3 snail A 6 I-S 11197 1-3 bivalve A 10 I-S 11198 1-4 bivalve M 9 1-S 11198 1-4 Costatoria sp. 8 1-S 11198 1-4 Halobia sp. 2 I-S 11198 1-4 indet. evolute ammonite 1 I-S 11198 1-4 indet. involute ammonite 10 I-S 11198 1-4 indet. nautiloid 13 I-S 11198 1-4 Isocrinus sp. 1 I-S 11198 1-4 Klamathites sp. 6 1-S 11198 1-4 Lopha montis-caprilis 112 1-S 11198 1-4 Michelinoceras sp. 4 1-S 11198 1-4 Myophoria shoshoniensis 3 1-S 11198 1-4 Nuculana sp. 3 1-S 11198 1-4 Plicatula sp. 1 1-S 11198 1-4 Septocardia sp. 856 I-S 11198 1-4 snail A 160 I-S 11198 1-4 snail T 3 1-S 11198 1-4 cf. Nuculana 5 1-S 11199 1-5 Germanonautilus kummeli 1 1-S 11199 1-5 indet. ammonite 9 1-S 11199 1-5 Juvavites sp. 4 1-S 11199 1-5 Lopha montis-capriiis 4 1-S 11199 1-5 Michelinoceras sp. 3 1-S 11199 1-5 Myophoria shoshoniensis 1 1-S 11199 1-5 Septocardia sp. 24 1-S 11199 1-5 snail A 24 1-S 11199 1-5 270

TABLE C.1. (continued)

Taxon Number of Biofacies- Locality Sample Individuals Community Number Number Germanonautilus kummeli 3 1-S 11200 1-6 indet. ammonite 4 1-S 11200 1-6 Myophoria shoshoniensis 1 l-S 11200 1-6 Nuculana sp. 8 1-S 11200 1-6 Septocardia sp. 18 1-S 11200 1-6 snail A 28 1-S 11200 1-6 cf. Nuculana 14 1-S 11201 1-7 Germanonautilus kummeli 1 1-S II20I 1-7 indet. ammonite 16 1-S 11201 1-7 Klamathites sp. I 1-S 11201 1-7 Lopha montis-caprilis 8 1-S 11201 1-7 Pinna sp. 1 1-S 11201 1-7 Septocardia sp. 73 1-S 11201 1-7 snail A 50 1-S 11201 1-7 cf. Nuculana 24 1-S 11202 1-8 conc biv. 1 1-S 11202 1-8 Septocardia sp. 25 1-S 11202 1-8 snail A 74 1-S 11202 1-8 bivalve A 8 1-S 11203 1-9 Myophoria shoshoniensis 1 1-S 11203 1-9 Nuculana sp. 11 1-S 11203 1-9 Septocardia sp. 3 1-S 11203 1-9 snail A 3 1-S 11203 1-9 bivalve A 24 1-S 11204 l-IO indet. nautiloid 2 1-S 11204 l-IO Michelinoceras sp. 1 1-S 11204 1-10 Nuculana sp. 15 1-S 11204 1-10 Septocardia sp. 40 1-S 11204 1-10 snail A 8 1-S 11204 MO bivalve A 1 1-S 11205 l-ll bivalve M 4 1-S 11205 1-11 indet. brachiopod 1 1-S 11205 1-11 Lopha montis-caprilis 1 1-S 11205 1-11 Myophoria shoshoniensis 1 1-S 11205 1-11 Septocardia sp. 86 1-S 11205 1-11 snail A 1 1-S 11205 l-ll Arcavicula sp. 1 2-N 11206 1-12 bivalve A 11 2-N 11206 1-12 indet. bivalve 18 2-N 11206 1-12 indet evolute ammonite 3 2-N 11206 1-12 indet. involute ammonite 3 2-N 11206 1-12 Klamathites macrolobatus 4 2-N 11206 1-12 Nuculana {Thestyledd) sp. 1 2-N 11206 1-12 snail A 7 2-N II206 1-12 indet. bivalve 8 1-S 11207 1-13 Klamathites sp. 1 1-S 11207 1-13 Lopha montis-caprilis 2 1-S 11207 1-13 Septocardia sp. 29 1-S 11207 1-13 snail A 5 1-S 11207 1-13 271

TABLE C.I. (continued)

Taxon Number of Biofacies- Locality Sample Individuals Community Number Number Tropites sp. I 1-S 11207 1-13 Arcavicula sp. I 2-T 11208 1-14 Halobia sp. 14 2-T 11208 1-14 indet. bivalve 60 2-T 11208 1-14 indet. evolute ammonite 7 2-T 11208 1-14 indet. involute ammonite 6 2-T 11208 1-14 indet. nautiloid I 2-T 11208 1-14 Klamathites macrolobattis I 2-T 11208 1-14 Klamathites sp. 2 2-T 11208 1-14 Septocardia sp. I 2-T 11208 1-14 snail A 1 2-T 11208 1-14 Tropites crassicostatus 4 2-T 11208 1-14 Tropites latiumbilicatiis 2 2-T 11208 1-14 Tropites sp. 112 2-T 11208 1-14 Tropites subquadratus 2 2-T 11208 I-I4 cf. Nuculana 1 2-T 11209 1-15 indet. bivalve 3 2-T 11209 1-15 indet. ammonite 2 2-T 11209 1-15 Septocardia sp. I 2-T 11209 1-15 Tropites sp. 17 2-T 11209 1-15 Halobia sp. 3 2-N 11210 1-16 indet. ammonite 38 2-N 11210 1-16 MicheUnoceras sp. 2 2-N 11210 1-16 Nuculana sp. 30 2-N 11210 1-16 Tropites sp. 17 2-N 11210 1-16 ?juvenile ammonite 9 2-M 11211 1-17 cf. Guembelites sp. 26 2-M 11211 1-17 cf. Mojsisovicsites sp. 42 2-M 11211 1-17 cf. Nuculana 2 2-M 11211 1-17 Tropiceltites columbianus 16 2-M 11211 1-17 cf. Guembelites sp. 2 2-H 11212 1-18 Halobia sp. 5 2-H 11212 1-18 ?juvenile ammonite 39 2-M 11213 1-19 Arcestes sp. 12 2-M 11213 1-19 cf. Guembelites sp. 59 2-M 11213 1-19 cf. Mojsisovicsites sp. 43 2-M 11213 1-19 cf Nuculana 3 2-M 11213 1-19 Guembelites jandianus 2 2-M 11213 1-19 Guembelites sp. 8 2-M 11213 1-19 Halobia sp. 22 2-M 11213 1-19 indet. echinoid I 2-M 11213 1-19 Klamathites sp. 3 2-M 11213 1-19 MicheUnoceras sp. 3 2-M 11213 1-19 Mojsisovicsites sp. 19 2-M 11213 1-19 Tropiceltites columbianus 24 2-M 11213 1-19 Tropites sp. 1 2-M 11213 1-19 Halobia sp. 42 2-H 11214 1-20 Arcestes sp. 2 2-M 11215 1-21 cf. Guembelites sp. 21 2-M 11215 1-21 272

TABLE C.I. (continued)

Taxon Number of Biofacies- Locality Sample Individuals Community Number Number cf. Mojsisovicsites sp. 14 2-M I121S 1-21 Guembelites jandianus 1 2-M 11215 1-21 indet. bivalve 3 2-M 11215 1-21 Michelinoceras sp. I 2-M 11215 1-21 Tropiceltites columbianus I 2-M 11215 1-21 cf. Nuculana 1 3-na 11216 1-22 Halobia sp. 1 3-na 1I2I6 1-22 indet. ecliinoid I 3-na II2I6 1-22 indet. bivalve II 3-na 11216 1-22 indet. colonial coral 6 3-na 11216 1-22 indet. solitary coral I 3-na II216 1-22 indet. ammonite I 3-na 11216 1-22 Plectoconcha aequiplicata 2 3-na 11216 1-22 Septocardia sp. 2 3-na 11216 1-22 snail A I 3-na 11216 1-22 Zeilleria sp. 17 3-na 11216 1-22 indet. bivalve 1 3-P 11217 1-23 Lopha montis-caprilis 2 3-P 11217 1-23 Plectoconcha aequiplicata 39 3-P 11217 1-23 snail A 2 3-P 11217 1-23 Zeilleria cf. elliptica 10 3-P II217 1-23 Zugmayerella sp. 3 3-P 11217 1-23 Plectoconcha aequiplicata 12 3-P 11218 1-24 indet. brach. 3 3-P 11219 1-25 Plectoconcha aequiplicata 126 3-P 11219 1-25 snail A 3 3-P II219 1-25 Zugmayerella sp. 2 3-P I12I9 1-25 Plectoconcha aequiplicata 6 3-P 11220 1-26 Septocardia sp. I 3-P 11220 1-26 snail A 3 3-P 11220 1-26 Zeilleria cf elliptica 1 3-P 11220 1-26 Arcavicula sp. 1 3-P 11221 1-27 bivalve A I 3-P 11221 1-27 indet. bivalve 3 3-P 11221 1-27 Lopha montis-caprilis 16 3-P 11221 1-27 Myophoria shoshoniensis 1 3-P 11221 1-27 Plectoconcha aequiplicata 41 3-P 11221 1-27 snai! A 2 3-P 11221 1-27 snail T 3 3-P 11221 1-27 Arcavicula sp. 1 3-P 11222 1-28 indet. bivalve 11 3-P 11222 1-28 indet. colonial coral 1 3-P 11222 1-28 Plectoconcha aequiplicata 105 3-P 11222 1-28 snail A 4 3-P 11222 1-28 snail T 6 3-P 11222 1-28 Zeilleria cf. elliptica 1 3-P 11222 1-28 bivalve A 8 3-P 11223 1-29 Halobia sp. 5 3-P 11223 1-29 indet. bivalve 10 3-P 11223 1-29 273

TABLE C.1. (continued)

Taxon Number of Biofacies- Locality Sample Individuals Community Number Number Lopha montis-caprilis 48 3-P 11223 1-29 Plectoconcha aequiplicata 93 3-P 11223 1-29 snail A 12 3-P 11223 1-29 snail T 2 3-P 11223 1-29 Cinnabaria expansa 4 3-Z 11224 1-30 indet. echinoid 1 3-Z 11224 1-30 indet. bivalve 5 3-Z 11224 1-30 indet. colonial coral 1 3-Z 11224 1-30 indet. ammonite 1 3-Z 11224 1-30 Lopha montis-caprilis 55 3-Z 11224 1-30 Nevadalithamia cylindrica 2 3-Z 11224 1-30 Pinna sp. 1 3-Z 11224 1-30 Septocardia sp. 9 3-Z 11224 1-30 snail A 58 3-Z II224 1-30 snail T 24 3-Z 11224 1-30 Zeiiieria cf. eiliptica 72 3-Z 11224 1-30 indet. bivalve 3 3-Z 11225 1-31 Lopha montis-caprilis 8 3-Z 11225 1-31 Septocardia sp. 4 3-Z 11225 1-31 snail A 32 3-Z 11225 1-31 snail T 3 3-Z 11225 1-31 Zeiiieria cf. eiliptica 48 3-Z 11225 1-31 Lopha montis-caprilis 13 3-Z II226 1-32 snail A 2 3-Z 11226 1-32 snail T 5 3-Z 11226 1-32 Zeiiieria cf eiliptica 22 3-Z 11226 1-32 Isocrinus sp. 2 3-Z 11227 1-33 Lopha montis-caprilis 17 3-Z 11227 1-33 snail A 4 3-Z 11227 1-33 snail T 5 3-Z 11227 1-33 Zeiiieria cf eiliptica 36 3-Z 11227 1-33

*Key to Biofacies-Community: Communities were assigned to biofacies 1, 2, or 3 by Hogier (1992a). Community (Hogier, 1992a): L=Lopha S=Septocardia U=Nuculana T=Tropites H=Halobia M=MoJsisovicsites-Guembelites P=Plectoconcha Z=Zeilleria na=not assigned to a community 274

TABLE C.2. Number of individuals, taxa, biofacies, and communities listed in Section I of Hogler (1992a) and grouped by lithofacies (this dissertation). Life habit assignments are from Hogler (1992a) except for Halobia sp., which has my interpretation. Life habit key: p=pelagic, e=epifaunal, i=infaunal; sc=scavenger, cn=camivore, su=suspension-feeder, dp=deposit feeder. Table C.l lists abundance by sample from Hogler (1992a).

Biofacies-Community / Lithofacies Taxon Life I-L l-S 2-T 2-N 2-H 2-M 3-na 3-P 3-Z Abundance in habit Sla Sla Sbl Sbl Scs Sbl- n/a Pbl-O F Section I Pel ?juvenile ainmonite p/sc/cn 48 48 Arcestes sp. p/sc/cn 14 14 cf. Guembeliles sp. p/sc/cn 2 106 108 Guembeliies jandianus p/sc/cn 3 3 Guembeliles sp. p/sc/cn 8 8 indet. evolute ammonite p/sc/cn 1 7 3 11 indet. involute ammonite p/sc/cn 10 6 3 19 indet. ammonite p/sc/cn 29 2 38 1 1 71 Juvavites sp. p/sc/cn 4 4 Klamathites macrolobatus p/sc/cn 1 4 5 Klamathites sp. p/sc/cn 8 2 3 13 cf. Xfojsisovicsites sp. p/sc/cn 99 99 Mojsisovicsites sp. p/sc/cn 19 19 Tropiceitites columbianus p/sc/cn 41 41 Tropiies crassicostatus p/sc/cn 4 4 Tropites latiumbilicatus p/sc/cn 2 2 Tropiies sp. p/sc/cn 1 129 17 1 148 Tropites subquadratus p/sc/cn 2 2 Ammonoids—total 0 53 155 65 2 342 1 0 1 619

Germanonautilus kummeli p/sc/cn 5 5 indet. nautiloid p/sc/cn 1 19 1 21 Michelinoceras sp. p/sc/cn 8 2 4 14 Nautiloids—total I 32 I 2 0 4 0 0 0 40

Arcavicula sp. e/su 1 I 2 4 Lopha montis-caprilis e/su 59 150 66 93 368 Pinna sp. e/su 1 1 2 Plicatula sp. e/su 1 1 Bivalves E7SU—total 59 152 1 1 0 0 0 68 94 375 cf. Nuculana i/dp 43 1 5 1 50 Nucuiana (Thestyieda) sp. i/dp 1 I Nuculana sp. i/dp 37 30 67 Bivalves I/DP—total 0 80 1 31 0 5 1 0 0 118

Costatoria sp. i/su 2 8 10 Myophoria shosfioniensis i/su 7 1 8 Septocardia sp. i/su 53 1355 2 2 1 13 1426 Bivalves I/SU—total 55 1370 2 0 0 0 2 2 13 1444 275

TABLE C.2. (continued) Biofacies-Community / Lithofacies Taxon Life l-L I-S 2-T 2-N 2-H 2-M 3-n/a 3-P 3-Z Abundance in habit Sla Sla Sbl Sbl Scs Sbl- n/a Pbl-O F Section 1 Pel Halobia sp. p/su 4 2 14 3 47 22 1 5 98 Bivalves P/SU—total 4 2 14 3 47 22 1 5 98

Plectoconcha aequiplicata e/su 2 422 424 Zeilleria cf. elliptica e/su 12 178 190 Zeilleria sp. e/su 17 17 Zugmayerella sp. e/su 1 5 6 indet. brachiopod e/su 1 3 4 Brachiopods—total I 1 0 0 0 0 19 442 178 641 snail A 21 359 1 7 1 26 96 511 snail T 3 11 37 51 snail W 8 8 Gastropods—total 29 362 1 7 0 0 1 37 133 570

Taxa In Section 1 but not included in Figure FA: Cinnabaria expansa e/su 4 4 Nevadalithamia cylindrica e/su 2 2 indet. colonial coral e/su 2 6 1 1 10 indet. solitary coral e/su 1 1 indet. echinoid 1 1 1 3 Isocrinus sp. e/su 1 2 3 decapod sc/cn 1 1 bivalve A 43 11 9 63 bivalve M 13 13 conc biv. 1 1 indet. bivalve 8 63 18 3 11 25 8 136 Abundance by facies 151 2119 238 138 49 377 44 589 437 Total number of individuals reponed from Section I (Hogler, 1992a): 4142 276

TABLE C.3. Number of individuals by lithofacies, used in Figure 2.16. See also Tables C.I and C.2. Data from locality 11216 of Hogler (1992a; biofacies 3-na) are not used in this table.

LITHOFACIES Number of Sla Sbl Scs Sbl-Pel Pbl-O F Taxa Ammonoids 11 + 53 220 2 342 0 1 Nautiloids 2+ 33 3 0 4 0 0 Bivalve i/su 3 1425 2 0 0 2 13 Bivalve e/su 4 211 2 0 0 68 94 Bivalve p/su 1 6 17 47 22 5 0 Bivalve i/dp 1 + 80 32 0 5 0 0 Brachiopods 3+ 2 0 0 0 442 178 Gastropods 3+ 391 8 0 0 37 133 277

APPENDIX D

MARINE REPTILE DATABASE TABLE RELATIONSHIPS, STRUCTURES, AND FIELD CODES 278

LOCALITIES TABLE

LOC ID

MMR_TAXA TABLE Loc_ID MMRJD Lith ID Loc [D

LITHOLOGiES TABLE

LITH ID

FIGLFRE D.I. Marine reptile database table relations. Each box represents a table and arrows represent links among tables via key fields. Primary key fields are shown in capitals, foreign keys are shown in italics, and composite primary keys are shown in normal type. The rounded box represents a linking table and contains a composite primary key (Loc ID and Lith ID fields). In a relational database, a table describes records in terms of a collection of fields of a particular subject. The primary key field uniquely identifies each record in a table (Hernandez, 1997). 279

TABLE D. I. Marine reptile database table structures, including fields and explanations. Structures were developed with reference to Damuth et al. (1997), Hernandez (1997), T.L. Moore (personal communication, 1997), and Ziegler et al. (1985). Field names preceded by an asterisk are shown in Tables G.I-G.2. Unmarked fields were either redundant with the information shown in Tables G.I-G.2 or not consistently reported in the literature. Data not shown in Tables G. I-G.2 are available on request from the author. TABLE FIELD EXPLANATION LOCALITIES •LocJD Primary key for Localities table; uniquely identifies each locality Country Describes location State Describes location County Describes location Locality Describes location Lat_min_deg Describes location with geographical coordinates Lonmindeg Describes location with geographical coordinates *LatLonErrt Estimated error of geographical coordinates * Latdec Latitude in decimal degrees * Londec Longitude in decimal degrees Recon_lat Paleolatitude of locality using reconstructed paleogeography Reconjon Paleolongitude of locality using reconstructed paleogeography System Describes age Series Describes age Stage Describes age Substage Describes age * Lowerbound Maximum age of the locality, to stage if possible *Upper_bound Minimum age of the locality, to stage if possible *Stage_res Stage-restricted time interval code (see Table 3.2) *Stage_un Stage-unrestricted time interval code (see Table 3.2) •Series_un Series-unrestricted time interval code (see Table 3.2) ChronoAge Name of the chronostratigraphic unit the locality is assigned to ChronoType Kind of organism that names the chronostratigraphic unit *Age_Relt Estimated error of die assigned age * Formation Describes the lithologic unit * Member Describes the lithologic unit Thickness Describes the lithologic unit (in meters) *Strat_prect Estimated error of placement of the fossil locality * Lith_relt Estimated quality of the reporting of lithological data Assemblage Summary of associated fauna at the locality * Locref Sources of data Loc_page Page numbers of the reference(s) where data were obtained Loc memo Additional information about the locality tThese 4 fields were combined into a Reliability code in Table F.I. See Table D2 for explanation of codes. 280

TABLE D.I (continued) TABLE FIELD EXPLANATION

MMR TAXA •MMR-ID Primary key for the MMR Taxa table; uniquely identifies each record •LocJD Foreign key for the MMR Taxa table; links MMR Taxa and Localities tables OrigOrder Order reported »MMR_Order Order modified if necessary by taxonomic scheme (Appendix E) OrigFamily Family reported •MMRFamily Family modified if necessary by taxonomic scheme (Appendix E) OrigGenus Genus reported •MMR_Genus Genus modified if necessary by taxonomic scheme (Appendix E) Orig^Sp Species reported •MMR_Species Species modified if necessary by taxonomic scheme (Appendix E) Orig_abun Number of individuals reported from the locality Approxabun Generalized abundance rating •Complete Skeletal elements reported Non_skel Non-skeletal evidence reported (e.g., gastroliths, stomach contents) • Preservatn Estimate of degree of preservation and articulation MMR_size Generalized size rating of an animal »MMR_ref Sources of data MMR_page Page numbers of the reference(s) where data were obtained MMR memo Additional information about the marine reptile fossils 281

TABLE D.2. Database field codes for selected fields shown in Table G.I. Developed with reference to Damuth et al. (1997), Hernandez (1997), T.L. Moore (personal communication, 1997), and unpublished materials compiled by A.M. Ziegler ("Standard Procedures for Paleogeographic Data Compilation and Interpretation", 1975^ FIELD CODES Loc ID Attaches a record number to each datapoint. The 2-letter prefix of the number groups datapoints according to continent in the following format: "prefix"-"number" (e.g., AS-657 for a locality in China). Continent codes: AF Africa AN Antarctica AS Asia AU Australia and New Zealand EU Europe NA North America SA South and Central America

Map Interval 3-part code that describes the age resolution of the record. Form of the code is x-x-x. Character Meaning See Table 3.2 for criteria used to assign map 1 Stage res interval codes. 2 Stageun 3 Series un

Reliability 4-part code that describes the spatial and stratigraphic resolution of the record. Form of the code is xxx-x-#-xxx. A "$" indicates a null value, used where information was not reported in the literature. Character Meaning 1-3 (first set) LatLonErr (see below) 4 (second set) Age rel (see below) 5 (third set) Stratjjrec (see below) 6-8 (fourth set) Lithrel (see below)

LatLonErr Basic rating: A - Position reported in lat-lon coordinates B - Position plotted on map, estimated error < 0.5° C - Position plotted on map, estimated error < 1.0° D - Position plotted on map, estimated error > 1.0° Error rating modifiers: E - Locality name reported, coordinates from atlas G - Locality name reported, coordinates from GeoNet Names Server * - Coordinates fi'om nearest city (usually reported in the reference)

AgeRel A - well-dated to substage B - well-dated to stage C - well-dated to series D - guess or poorly resolved 282

TABLE D.2 (continued) FIELD CODES Reliability Stratj)rec (continued) I - specific bed 2 - specific member of a formation 3 - specific formation 4 - specific locality, formation not specified but local biozone specified 5 - specific locality, formation not specified but regional or global biozone specified 6 - specific locality, formation and biozone not specified 7 - general area (e.g., a mountain range)

Lith_rel D - stratigraphic column E - geologic map F - locality or quarry map 283

TABLE D.3. Minutes to decimal degrees conversions.

Minutes Decimal degrees 58-03 0.0 04-09 0.1 10-15 0.2 16-21 0.3 22-27 0.4 28-33 0.5 34-39 0.6 40-45 0.7 46-51 0.8 52-57 0.9 284

TABLE D.4. Database field codes for selected fields shown in Table G.2. Developed with reference to Damuth et al. (1997), Hernandez (1997), T.L. Moore (personal communication, 1997), and unpublished materials compiled by A.M. Ziegler ("Standard Procedures for Paleogeographic Data Compilation and Interpretation", 1975^ FIELD CODES LocID See Table D2 Map Interval See Table D.2 MMR ID MR-####. The 2-letter prefix indicates that the record contains taxonomic information for one taxon at one locality.

Complete A multi-value field. All elements reported from the locality were recorded. a - whole skeleton b - skull c - mandibles d - teeth e - vertebrae f - ribs g - gastralia h - girdle i - limb j - dermal bone k - carapace q - composite, all major elements from >1 specimen z - fragments or unspecified bones $ - not reported

Preservatn 1 - isolated teeth are the only consistently identifiable elements 2 - robust fragments preserved - jaw fragments, teeth, ends of long bones 3 - intermediate between robust and delicate fragments 4 - fi^gmentary material, but delicate structures are preserved 3 - parts disarticulated but largely whole; e.g., bone beds, individuals cannot be recognized 6 - parts largely disarticulated, but individuals can often be recognized 7 - occurrences mostly or completely articulated 0 - not reported 285

APPENDIX E

TAXONOMIC SCHEMES USED IN THE MARINE REPTILE DATABASE 286

TABLE E. 1. Taxonomic scheme for the Ichthyosauria. Compiled from Bardet 1995; Fernandez 1994, 1997; McGowan 1994; McGowan 1995a, 1995b, 1995c; McGowan 1996a, 1996b; McGowan 1997; Motani 1997; Nicholls and Brinkman 1995; and Sander 1997. Age ranges (stage) from Bardet 1995. Taxa whose position in the scheme is uncertain (e.g., taxa in the midst of revision) are listed in the last part of the table. Order Family Genus Species Age range Ichthyosauria incertae sedis Chacaicosaurus Fernandez 1994 BAJ cayi Fernandez 1994 Baj Chaohusaurus Young & Dong 1972 SPA geishanensis Young & Dong 1972 Spa Chensaunis Mazin et al. 1991 SPA chaoaxianensis (Chen 1985) Spa faciles (Chen 1985) Spa Grippia Wiman 1928 SPA longirostris Wiman 1928 Spa Himalayasaurus Dong 1972 NOR tibetensis Dong 1972 Nor Hudsonelpiclia McGowan 1995 NOR brevirostris McGowan 1995 Nor Parvinatator Nicholls & Brinkman 1995 7ANS-LAD waipitiensis Nicholls & Brinkman 1995 Ans-Lad Pessosaurus Wiman 1910 ANS-LAD polaris (Hulke 1873) Lad Phalarodon Merriam 1910 ANS-LAD fraasi Merriam 1910 Ans-Lad Svalbardosaurus Mazin 1981 SPA crassidens Mazin 1981 Spa Thaisaurus Mazin etal. 1991 SPA chonglakmanii Mazin etal. 1991 Spa Tibetosaurus Wang 1982 NOR tingjiensis Wang 1982 Nor Utatsusaurus Shikamaetal. 1978 SPA hataii Shikamaetal. 1978 Spa Mixosauridae MLxosaurus Baur 1887 SPA7-LAD atavus (Quenstedt 1852) Ans comalianus (Bassani 1886) Ans/Lad maotaiensis Young 1965 Ans nutans (Merriam 1908) Ans-Lad nordenskioeldi (Hulke 1873) Spa?-Lad Shastasauridae Califomosaurus Kuhn 1934 CRN perrini (Merriam 1902) Cm 287

TABLE E.l. (continued) Order Family Genus Species Age range Cymbospondylus Leidy 1868 SMI-LAD petrinus Leidy 1868 Ans buchseri Sander 1989 Ans/Lad nevadanus Merriam 1908 Ans piscostis Leidy 1868 Ans Merriamia Boulenger 1904 CRN zitteli (Merriam 1903) Cm Shastasaurus Merriam 1895 ANS7-NOR pacificus Merriam 1895 Cm neoscapularis McGowan 1994 Nor neubigi Sander 1997 Ans Toretocnemus Merriam 1902 CRN califomicus Merriam 1902 Cm Shonisauridae Shonisaurus Camp 1976 CRN-NOR popularis Camp 1976 Cm Ichthyosauridae Baptanodon Marsh 1880 OXF nutans (Marsh 1879) Oxf discus (Marsh 1880) Oxf marshi Knight 1903 Oxf reedi Gilmore 1907 Oxf robustus Gilmore 1906 Oxf Ichthyosaurus De la Beche & Conybeare 1821 NOR-SIN communis Conybeare 1822 Het-Sin breviceps Owen 1881 Het-Sin conybeari Lydekker 1888 Het-Sin Janiceps McGowan 1996 Nor Ophthalmosaurus Seeley 1874 CLV-TTH icenicus Seeley 1874 Civ monocharactus Appleby 1956 Clv-Tth Brachyptergius Huene 1922 KIM-TTH extremus (Boulenger 1904) Tth Kuhn 1934 Caypullisaurus Fernandez 1997 TTH bonapartei Femandez 1997 Tth Stenopterygidae Stenopterygius Jaekel 1904 TOA quadriscissus (Quenstedt 1858) Toa cuneiceps McGowan 1979 Toa hauffianus Huene 1922 Toa longipes (Wurstemberger 1876) Toa macrophasma McGowan 1979 Toa megacephalus Huene 1922 Toa 288

TABLE E. 1 • (continued) Order Family Genus Species Age range megalorhinus Huene 1922 Toa Leptopterygidae Eurhinosaurus Abel 1909 HET-TOA longirostris (Mantell 1851) Het-Toa Excalibosaurus McGowan 1986 SIN costini McGowan 1986 Sin Temnodontosauridae Leptonectes McGowan 1996 RHE-SIN tenuirostris (Conybeare 1822) Rhe-Plb solei McGowan 1993 Sin

Temnodontosaurus Lydekker1889 HET-SIN platyodon (Conybeare 1822) Het-Sin eurycephalus McGowan 1974 Sin burgundiae (Gaudry 1892) Toa Playtpterygidae Nannopterygius Huene 1922 FOM enthekiodon (Hulke 1870) Kim Platypterygius Huene 1922 APT7-CEN "campylodon" (Carter 1846) Alb-Cen americanus (Nace 1939) Alb-Cen hercynicus (Kuhn 1946) Apt kiprijanojfi (Romer 1968) Alb-Cen longmani Wade 1990 Alb plalydactylus Broili 1907 Apt

Taxonomic position uncertain or pending; ?Simbirsldasaurus birjukovi Otschev & Efimov 1985 Hau ?"Platypterygius " hauthali (Huene 1927) Ber Ichthyosaurus votgensis Ichthyosaurus queenstedti? Leptopterygius disinteger Stenopterygius acutirostris Stenopterygius longifrons 289

TABLE E.2. Taxonomic scheme for the Plesiosauria. Compiled from Adams 1997; Bakker 1993; Bardet 1995; Bardet & Godefroit 1998; Blodgettet al. 1995; Carpenter 1996, 1997; Gasparini & Goni 1985; Godefroit 1994; Hampe 1992; Motani 1997; Otschev 1976, 1977; Sander et al. 1997; Storrs 1997; and Storrs & Taylor 1996. Age ranges (stage) fi-om Bardet 1995. Taxa whose position in the scheme is uncertain are listed last. Order Superfamily Family Genus Species Age Range Sauropterygia incertae sedis Omphalosauridae Omphalosaurus Merriam 1906 SPA-ANS nevadanus Merriam 1906 Ans nettarhynchus Mazin & Bucher 1987 Spa Pleiosauria nisseri (Wiman 1910) Ans Pistosauroidea Pistosauridae Augustasaurus Sander etal. 1997 hagdomi Sander etal. 1997 Pistosaurus Meyer 1839 ANS longaevus Meyer 1839 Ans Plesiosauroidea incertae sedis Sthenarosaurus Watson 1909 TOA dawkinsi Watson 1909 Toa Thalassiodracon Storrs & Taylor 1996 hawkinsi (Owen 1840) Plesiosauridae Plesiosaurus Conybeare 1821 HET-TOA dolichodeirus Conybeare 1824 Het guilelmiimperatoris Dames 1893 Toa "P." toumemirensis Sciauetal. 1990 Toa Attenborosaurus Bakker 1993 SIN conybeari Sollas 1881 Sin Elasmosauridae Alzadasaurus Welles 1943 APT-CMP riggsi Welles 1943 Tur colombiensis Welles 1962 Apt kansasensis Welles 1952 San-Cmp pembertoni Welles & Bump 1949 Cmp Aphrosaums WeUes 1943 MAA fitrlongi Welles 1943 Maa Aristonectes Cabrera 1941 MAA parvidens Cabrera 1941 Maa Brancasaurus Wegner 1914 BER brancai Wegner 1914 Ber Elasmosaurus Cope 1868 TUR-CMP platyurus Cope 1868 Cmp Eretmosaurus Seeley 1874 SIN rugosus (Owen 1840) Sin 290

TABLE E.2. (continued) Order Superfamily Family Genus Species Age Range Fresnosaurus Welles 1943 MAA drescheri Welles 1943 Maa Hydralmosaunis Welles 1943 SAN-CMP serpentinus (Cope 1877) San-Cmp Hydrotherosaurus Welles 1943 MAA alexandrae Welles 1943 Maa Libonectes Carpenter 1997 morgani (Welles 1949) Leurospondylus Brown 1913 MAA ultimus Brown 1913 Maa Mauisaurus Hector 1874 MAA haasti Hector 1874 Maa Microcleidus Watson 1909 TOA hotnalospondylus (Owen 1865) Toa macropterus (Seeley 1865) Toa Morenosaurus Welles 1943 MAA stocki Welles 1943 Maa Mortumeria Chatteijee & Creisler 1994 MAA seymourensis (Chatteijee& Small 1989) Maa Muraenosaurus Seeley 1874 CLV leedsii Seeley 1874 Civ beloclis Seeley 1892 Civ Slyxosaurus Welles 1943 CMP browni Welles 1943 Cmp Thalassomedon Welles 1943 CEN haningtoni Welles 1943 Cen new sp. Storrs & Langston (in press) Cen Tuarangisaurus Wiffen & Moisley 1986 CMP-MAA keyesi Wiffen & Moisley 1986 Cmp-Maa Cryptoclididae Colymbosaurus Seeley 1874 KJM-TTH trochanterius (Owen 1840) Kim-Tth Cryptoclidus Seeley 1892 CLV eurymerus (Phillips 1871) Civ richardsoni (Lydekker 1889) Civ Tricleidus Andrews 1909 CLV seeleyi Andrews 1909 Civ Pliosauroidea Pliosauridae Archaeonectrus Novozhilov 1964 SIN rostratus (Owen 1865) Sin Bishanopliosaurus Dong 1980 TOA youngi Dong 1980 Toa Brachauchenius Williston 1903 CEN7-TUR 291

TABLE E.2. (continued) Order Superfamily Family Genus Species Age Range lucasi Williston 1903 Cen?-Tur Eurycleidus Andrews 1922 HET arcuatus (Owen 1840) Het megacephalus (Smtchbury 1846) Het Kronosaurus Longman 1924 APT-ALB queenslandicus Longman 1924 Apt-Alb boyacensis Hampe 1992 Leptocleidus Andrews 1922 BRM superstes Andrews 1922 Brm Liopleurodon Sauvage 1873 CLV-TTH ferox Sauvage 1873 Civ macromerus (Phillips 1871) Kim-Tth pachydeirus (Seeley 1869) Civ rossicus (Novozhilov 1964) Tth Macroplata Swinton 1930 HET tenuiceps Swinton 1930 Het Megalneusaurus rex Knight 1898 Peloneustes Lydekker 1889 CLV philarchus (Seeley 1869) Civ Peyerus Stromer 1935 VLG capensis (Andrews 1911) Vlg Plesiopleurodon Carpenter 1996 CEN wellesi Carpenter 1996 Cen Pliosaurus Owen 1841 CLV-FGM brachydeirus Owen 1841 Kim andrewsi Tarlo 1960 Civ brachyspondylus (Owen 1839) Kim Polypiychodon Owen 1841 CEN-TUR interruptus Owen 1841 Cen hudsoni Welles & Slaughter 1963 Cen-Tur Rhomaleosaurus Seeley 1874 TOA zetlandicus Toa victor (Fraas 1910) Toa Simolestes Andrews 1909 CLV-TTH vorax Andrews 1909 Civ indicus (Lydekker 1877) Tth keileni Godefroit 1994 Baj ? Sinopliosaurus ? Jusuiensis Houetal., 1975 Strongylokrotaphus Novozhilov 1964 TTH irgisensis (Novozhilov 1948) Tth ? Yuzhoupliosaurus Zhang, 1985 ? chengjiangensis Zhang, 1985 292

TABLE E.2. (continued) Order Superfamily Family Genus Species Age Range Plesiosauroidea? Polycotylidae Dolichorhynchops Williston 1902 SAN-CMP osbomi Williston 1902 San-Cmp Polycotylus Cope 1869 CMP latipinnis Cope 1869 Cmp Trinacromerum Cragin 1888 ALB7-CMP? bentonianum Cragin 1888 Cen-Tur bonneri Adams 1997 kirki Russell 1935 Tur-Cmp? lafquenianum Gaspahni & Goni 198S willistoni Riggs 1944 Cen Georgiasaurus Otschev 1976 SAN

penzensis Otschev 1976 San Taxonomic position uncertain or pending: "Macroplata" longirostris (Blake 1876) "Plesiosaurus" propinquus Blake 1876 "Plesiosaurus" macrocephalus Owen 1840 "Muraenosaurus" mazetieri Bigot 1938 "Plesiosaurus" costatus Owen 1840 "Plesiosaurus" shirleyensis Knight 1900 Colymbosaurus sclerodirus Bogolubov 1911 Cryptocleidus simbirskensis Bogolubov 1909 Cryptocleidus aldingeri V. Huene 1935 Cryptocleidus? beaugrandi Sauvage 1912 Eretmosaurus dubius Blake 1876 Muraenosaurus fahrenkohli (Waldheim 1846) Pachycostasaurus dawni Cruickshank et al., 1996 Plesiosaurus ?subtrigonus Owen 1840 Plesiosaurus eleutheraxon Seeley Plesiosaurus trigonus Cuvier 1824 Plesiosaurus? baruthicus Kuhn 1934 Plesiosaurus? keuperinus V. Huene 1929 Plesiosaurus? pentagonus Cuvier 1824 Plesiosaurus? platydeirus Owen 1854 (non Seeley 1869) Plesiosaurus? robustus Dames 1895 Pliosaurus gamma Phillips 1871 Pliosaurus irgisensis (Novozhilov 1948) Pliosaurus rossicus Novozhilov 1948 Rhomaleosaurus? morincinus Sauvage 1879 Termatosaurus alberti Tb. Plieninger 1844 Tricleidus svalbardensis Persson 1962 Tricleidus? laramiensis (Knight 1900) 293

TABLE E.3. Taxonomic scheme for the family Mosasauridae. Compiled from Bardet 1995; Lingham- Soliar 1992, 1996, 1998; and Lingham-Soliar and Nolf 1989. Age ranges from Bardet 1995. Taxa whose position in the scheme is uncertain are listed last. Subfamily Genus Species Age Range incertae cedis Goryonosaurus Azzaroli et al. 1972 MAA nigeriensis (Swinton 1930) Maa Pluridens Lingham-Soliar 1998 walkeri Lingham-Soliar 1998 Mosasaurinae Amphekepubis Mehl 1930 SAN Johnsoni Mehl 1930 Carinodens Thurmond 1969 CMP-MAA belgiciis (Woodward 1891) Maa fraasi (Dollo 1913) Cmp-Maa Clidasies Cope 1868 CON-CMP iguanavus Cope 1868 Cmp liodontus Merriam 1894 Con-Cmp propython Cope 1869 San-Cmp Globidens Gilmore 1912 CMP-MAA alabamaensis Gimore 1912 Cmp dakotaensis Russell 1975 Cmp Leiodon Owen 1840 CMP-MAA anceps Owen 1840 Cmp-Maa compressidens Gaudry 1892 Cmp mosasauroides Gaudry 1892 Maa sectorius Cope 1871 Maa Mosasaurus Conybeare 1822 SAN-MAA hqffmanni Mantell 1829 Maa conodon (Cope 1881) Cmp-Maa dekayi Brown 1838 Maa flemingi Wiffen 1990 Cmp-Maa lemonnieri Dollo 1904 Maa lonzeensis Dollo 1904 San mangahouangae (Wiffen 1980) Cmp-Maa maximus Cope 1869 Maa missouriensis (Harlan 1834) Cmp-Maa mokoroa Gregg & Welles 1971 Maa Plotosaurus Camp 1951 MAA bennisoni (Camp 1942) Maa tuckeri (Camp 1942) Maa Rikisaurus Wiffen 1990 CMP-MAA tehoensis Wiffen 1990 Cmp-Maa Taniwhasaurus Hector 1874 MAA oweni Hector 1874 Maa Plioplatecarpinae Ectenosaurus Russell 1967 SAN-CMP clidastoides (Merriam 1894) San-Cmp Halisaurus Marsh 1869 SAN-MAA platyspondylus Marsh 1869 Maa 294

TABLE E.3. (continued) Subfamily Genus Species Age Range onchognathus (Merriam 1894) San-Cmp ortliebi (Dollo 1889) Maa Igdamanosaurus Lingham-Soliar 1991 CMP-MAA aegyptiacus (Zdansky 1935) Cmp-Maa Platecarpus Cope 1869 TUR-MAA? tympaniticus Cope 1869 San-Cmp bocagei (Antunes 1964) Tur coryphaeus (Cope 1872) Con ictericus (Cope 1871) San-Cmp somenensis Thevenin 1896 San-Cmp Plesiotylosaurus Camp 1942 MAA crassidens Camp 1942 Maa Plioplatecarpus Dollo 1882 CMP-MAA marshii Dollo 1882 Maa depressus (Cope 1869) Maa houzeaui Dollo 1889 Maa primaevus Russell 1967 Cmp ?crassartus Cope 1872 Cmp Prognathodon Dollo 1889 CMP-MAA solvayi Dollo 1889 Maa giganteus Dollo 1904 Maa overtoni (Williston 1897) Cmp-Maa rapax (Hay 1902) Maa waiparaensis Gregg & Welles 1971 Maa Selmasaurus Wright & Shannon 1988 CMP russelli Wright & Shannon 1988 Cmp Yaguarasaurus Paramo 1994 TUR columbianus Paramo 1994 Tur Tylosaurinae Williston 1897 Hainosaurus Dollo 1885 SAN-MAA bemardi Dollo 1885 Maa gaudryi (Thevenin 1896) San pembinensis NichoUs 1988 Cmp Tylosaurus Marsh 1872 CON-MAA proriger (Cope 1869) San-Cmp haumuriensis (Hector 1874) Maa nepaeolicus (Cope 1874) Con ? capensis Broom 1912 Upper K ? iembeensis (Telles-Antunes) Tur Taxonomic position uncertain or pending: Clidastes stembergi Wiman 1920 San-Cmp Mosasaurus camperi "Platecarpus" intermedius (Leidy 1870) Platecarpus ptychodon Arambourg 1952 Tylosaurus zangerli Russell 1970 295

APPENDIX F

PALEOGEOGRAPHIC BASE MAPS AND UPWELLING PREDICTIONS Figure F. i. Predicted upweiling (hachured areas) and paleogeography for the Anisian map interval (237.0 Ma). Darkest shading = highlands, medium shading = lowlands, and lightest shading = continental shelf. All subsequent maps follow this key. Predicted upweiling based on Parrish and Curtis (1982) and Parrish (1995); paleogeography from Scotese and Golonka (1992).

Figure F.2. Predicted upweiling and paleogeography for the Norian map interval (216.0 Ma). Figure F.3. Predicted upwelling and paleogeography for the Pliensbachian map interval (195.0 Ma).

Figure F.4. Predicted upwelling and paleogeography for the Callovian map interval (166.0 Ma). Figure F.S. Predicted upweiling and paleogeography for the Kimmeridgian map interval (152.2 Ma).

MM..

Figure F.6. Predicted upweiling and paleogeography for the Valanginian map interval (130.2 Ma). 299

Figure F.7. Predicted upweiling and paleogeography for the Aptian map interval (118.0 Ma).

Figure F.8. Predicted upweiling and paleogeography for the Cenomanian map interval (94.0 Ma). 300

Figure F.9. Predicted upwelling and paleogeography for the Coniacian map interval (88.0 Ma).

Figure F. 10. Predicted upwelling and paleogeography for the Maastrichtian map interval (69.4 Ma). 301

APPENDIX G

MARINE REPTILE DATABASE 302

TABLE G.l Marine Reptile Database Localities Age Range Lat Lon Maximum Minimum Iniw^ Reliability Reference

AF-239 28.2 29.0 Cenomanian Cenomanian h-h-h gd-B-S-S 229 AF-241 -33.8 25.6 Valanginian Hauterivian $-f-g b-b-3-S 229; 189 AF-314 25.1 32.8 Campanian Maastrichtian $-i-h d-b-3-f 90 AF-315 28.5 28.7 Cenomanian Cenomanian h-h-h d-b-3-f 90 AF-316 13.4 5.7 Maastrichtian Maastrichtian j-j-h e»-b-l-def 281 AF-317 13.4 5.7 Maastrichtian Maastrichtian j-j-h e«-b-l-def 281 AF-37 -8.3 13.3 Turonian Turonian $-i-h b-B-2-F 149 AF-38 -5.8 12.7 Maastrichtian Maastrichtian j-j-h egb-S-S-S 148 AF-39 -5.9 12.7 Maastrichtian Maastrichtian j-j-h egb-S-S-S 148 AF-398 -5.0 12.3 Maastrichtian Maastrichtian j-j-h b-b-3-f 293 AF-399 -7.2 12.9 Maastrichtian Maastrichtian j-j-h b-b-3-f 293 AF-400 -8.5 13.4 Maastrichtian Maastrichtian j-j-h b-b-3-f 293 AF-401 -14.3 12.4 Maastrichtian Maastrichtian j-j-h b-b-3-f 293 AF-402 -14.9 12.4 Maastrichtian Maastrichtian j-j-h a-b-3-f 293 AF-403 -15.1 12.2 Maastrichtian Maastrichtian j-j-h b-b-3-f 293 AF-404 -15.1 12.2 Maastrichtian Maastrichtian j-j-h b-b-3-f 293 AF-405 -5.3 12.4 Upper Cretaceous Upper Cretaccous $-$-h b-b-3-f 293 AF^06 -5.2 12.3 Upper Cretaceous Upper Cretaceous $-$-h b-b-3-f 293 AF-407 -5.0 12.2 Maastrichtian Maastrichtian j-j-h b-b-3-r 293 AF-408 -5.0 12.1 Maastrichtian Maastrichtian j-j-h b-b-3-f 293 AF-409 -12.6 13.4 Maastrichtian Maastrichtian j-j-h b-b-3-f 293 AF-410 -13.9 12.5 Maastrichtian Maastrichtian J-j-h b-b-3-f 293 AF-411 -14.3 12.4 Maastrichtian Maastrichtian j-j-h b-b-3-f 293 AF-412 -14.3 12.4 Maastrichtian Maastrichtian j-j-h a-b-3-f 293 AF-413 -14.7 12.2 Maastrichtian Maastrichtian j-j-h b-b-3-f 293 AF-414 -14.8 12.2 Maastrichtian Maastrichtian j-j-h b-b-3-f 293 AF^7 15.4 5.8 Maastrichtian Maastrichtian j-j-h a-b-3-$ 136; 146; 149 AF-544 31.6 -8.7 Maastrichtian Maastrichtian J-j-h b-b-4-cf 6 AF-545 32.9 -6.6 Maastrichtian Maastrichtian J-j-h b-b-4-ef 6 AF-546 32.5 -7.2 Maastrichtian Maastrichtian j-j-h b-b-4-ef 6 AF-548 32.6 •6.2 Maastrichtian Maastrichtian j-j-h b-b-«-ef 6 AF-549 32.3 -7.4 Maastrichtian Maastrichtian j-j-h b-b-4-ef 6 AF-550 32.3 -8.5 Maastrichtian Maastrichtian j-j-h b-b-4-ef 6 AF-55I 31.2 -8.9 Maastrichtian Maastrichtian j-j-h b-b-4-cf 6 AF-552 30.6 -9.0 Maastrichtian Maastrichtian j-j-h c-b-4-ef 6 AF-673 33.0 10.8 Anisian Ladinian $-a-b gb-c-6-S 120; 240 AF-691 -21.0 45.7 Campanian Campanian S-i-h gd-a-4-S 28 AF-732 33.2 10.3 Lower Cretaceous Lower Cretaceous $-$-g g-c-6-df 265 AF-760 -21.0 45.0 Portlandian Portlandian e-e-f gd-b-6-S 95 AF-761 26.0 27.0 Maastrichtian Maastrichtian j-j-h b-b-2-df 313 AF-83 -33.5 21.7 Valanginian Hauterivian $-f-g g*-b-3-S 126;189 AF-861 31.1 28.0 Campanian Campanian $-i-h gd-b-6-S 235 AF-862 29.5 31.6 Campanian Campanian S-i-h gd-b-6-S 235 AF-896 15.6 5.8 Maastrichtian Maastrichtian j-j-h a-b-3-f 152 AN-628 -64.3 -56.7 Campanian Maastrichtian S-i-h a-b-3-S 64; 62 AN-841 •63.9 -57.5 Campanian Campanian S-i-h b-b-6-S 107 AN-842 -64.3 -58.3 Campanian Campanian S-i-h e-b-6-S 107 AS-2 53.3 140.8 Middle Triassic Middle Triassic $-S-b gb-c-3-S 48 AS-212 52.5 44.9 Upper Cretaceous Upper Cretaceous S-S-h gc-C-S-S 229 AS-213 51.2 57.8 Upper Cretaceous Upper Cretaceous S-S-h gb-C-S-S 229 AS-214 51.5 45.5 Cenomanian? Cenomanian? h-h-h d-C-S-S 309;229 AS-215 51.7 39.2 Cenomanian Cenomanian h-h-h gd-B-S-S 229 AS-216 51.7 36.2 Cenomanian Cenomanian h-h-h gd-B-S-S 229; 244 AS-217 51.1 58.0 Upper Cretaceous Upper Cretaceous S-S-h gc-C-S-S 229 AS-218 55.8 35.6 Albian? Albian? S-h-g gd-C-S-S 229 AS-219 45.3 33.5 Lower Cretaceous Lower Cretaceous S-S-g d*-C-S-S 229 AS-220 57.8 41.2 Portlandian Portlandian e-e-f d*-B-S-S 229 AS-221 55.7 37.6 Portlandian Portlandian e-e-f ge-B-S-S 229 AS-222 55.8 37.5 Portlandian Portlandian e-e-f ge-B-S-S 229 303

TABLE G. 1 Marine Reptile Database Localities Age Range Lat Lot) Maximum Minimum Interv^ Reliability Reference

AS-223 53.8 37.4 Portlandian Portlandian e-e-f gc-B-S-S 229 AS-225 57.6 39.9 Callovian Oxfordian $-d-c gd-D-S-S 229 AS-226 54.6 39.7 Callovian Callovian d-d-e gd-B-S-S 229 AS-227 54,6 48.4 Callovian Callovian d-d-e gd-B-S-S 229 AS-228 59.2 39.9 Callovian? Callovian? d-d-e gd-C-S-S 229 AS-229 55.8 37.5 Portlandian? Portlandian? e-e-f ge-C-S-S 229 AS-230 53.8 37.4 Kimmcridgian Kimmeridgian $-e-f ge-B-S-S 229 AS-231 51.4 41.6 Kimmeridgian Kimmeridgian $-e-f g-B-S-S 229 AS-232 55.2 47.1 Kimmeridgian Kimmeridgian $-c-f gb-B-S-S 229 AS-233 57.6 43.5 Oxfordian Oxfordian S-e-f gd-B-S-S 229 AS-235 72.0 129.2 Lower Jurassic Lower Jurassic $-$-d ge-C-S-S 229 AS-236 47.2 142.5 Upper Cretaceous Upper Cretaceous $-S-h gd-C-S-S 229;244 AS-237 29.6 104.7 Oxfordian Oxfordian $-c-f gc-B-S-S 229 AS-238 32.6 34.9 Cenomanian Cenomanian h-h-h ge-B-S-S 229 AS-247 80.2 53.0 Oxfordian Oxfordian S-e-f gb-B-S-S 229 AS-291 48.6 43.2 Upper Cretaceous Upper Cretaceous $-S-h g-C-S-S 229; 244 AS-3 -9.8 124.5 Middle Triassic Middle Triassic $-S-b gb-c-6-S 48 AS-4 37.0 33.0 Middle Triassic Middle Triassic $-$-b gc-c-6-$ 48 AS-45 54.6 48.4 Kimmeridgian Kimmeridgian $-c-f g-A-2-$ 324 AS-49 51.7 36.1 Albian Cenomanian $-h-g e»-D-3-F 244; 17; 19 AS-586 34.5 135.0 Maastrichtian Maastrichtian j-j-h d-b-3-f 262 AS-587 69.3 143.5 Upper Triassic Upper Triassic $-$-c gc-c-6-S 215 AS-588 32.0 36.0 Maastrichtian Maastrichtian j-j-h d-b-3-df 8; 1 AS-593 10.8 78.7 Albian Cenomanian $-h-g gb-c-3-f 173; 19 AS-603 45.4 142.0 Cenomanian Cenomanian h-h-h b-b-2-f 214 AS-64 38.7 142.5 Spathian Spathian $-a-a gd*-C-3- 271;48 DEF AS-646 31.6 117.9 Spathian Spathian $-a-a gb-b-3-f 323; 20 AS-647 28.6 86.6 Norian Norian b-b-c gc-b-3-f 323; 87; 20 AS-65 39.6 142.0 Spathian Spathian S-a-a gd»-C-3- 271; 48 DEF AS-653 28.4 85.8 Norian Norian b-b-c gb-b-3-f 87; 48 AS-655 49.5 44.6 Upper Cretaceous Upper Cretaceous $-$-h gb-c-6-f 244 AS-656 49.2 37.3 Upper Cretaceous Upper Cretaceous $-$-h gb-c-6-f 244 AS-657 44.9 38.0 Senonian Senonian $-$-h gb-c-6-f 244 AS-658 49.1 57.2 Senonian Senonian S-$-h c-c-6-f 244 AS-659 52.9 63.0 Maastrichtian Maastrichtian j-j-h gc-b-6-f 244 AS-660 38.2 67.2 Lower Cretaceous Lower Crctaceous $-$-g e-c-6-f 244 AS-66I 40.9 45.9 Maastrichtian Maastrichtian j-j-h gb-b-6-f 244 AS-662 44.8 33.9 Maastrichtian Maastrichtian j-j-h gb-b-6-f 244 AS-67 27.9 106.3 Anisian Anisian J-a-b gc-C-6-S 325; 48 AS-677 8.2 98.8 Lower Triassic Lower Triassic $-S-a gb-c-5-S 172 AS-678 31.2 35.2 Campanian Campanian $-i-h gc-b-5-S 238 AS-688 23.6 70.0 Kimmeridgian Kimmeridgian S-e-f gc-a-2-f 24 AS-702 54.3 48.4 Hauterivian Hauterivian S-f-g gc-b-3-S 216 AS-706 42.9 142.1 Campanian Campanian S-i-h a-a-3-r 203 AS-708 29.9 106.4 Middle Jurassic Middle Jurassic S-S-e gc-c-6-S 328 AS-709 22.8 108 J Lower Cretaceous Lower Cretaceous S-$-g gc-c-6-df 128 AS-717 44.1 141.7 Santonian Santonian S-i-h c-a-3-S 165 AS-727 42.9 142.1 Maastrichtian Maastrichtian j-j-h d-b-3-S 213; 138 AS-728 43.2 141.9 Maastrichtian Maastrichtian j-j-h d-b-3-S 213; 138 AS-729 40.2 141.8 Santonian Santonian S-i-h d-b-3-S 213;138 AS-730 37.1 140.9 Santonian Santonian S-i-h d-b-3-S 213; 138 AS-738 59.7 150.2 Norian Norian b-b-c gc-b-6-$ 204; 48 AS-739 65.7 159.3 Camian Camian S-a-c gc-b-6-S 204; 48 AS-74I 62.0 160.4 Camian Camian S-a-c gc-b-6-S 48 AS-747 64.7 154.0 Ladinian Ladinian S-a-b gc-b-6-S 204; 48 AS-748 65.6 159 J Ladinian Ladinian S-a-b gd-b-6-S 204;48 AS-755 77.0 96.0 Lower Triassic Lower Triassic S-S-a gd-c-6-S 48 304

TABLE G.l Marine Reptile Database Localities ARC Range Lat Lon Maximum Minimum interim Reliability Reference

AS-765 62.0 120.4 Upper Lias Upper Lias $-c-d gd-c-6-S 204 AS-766 64.3 126.4 Toarcian Toarcian $-c-d gc-b-6-S 204 AS-767 63.8 121.6 Bajocian Bathonian $-d-e gd-c-6-$ 204 AS-770 63.2 48.0 Tithonian Tithonian e-e-f gb-b-6-S 204 AS-771 63.7 54.1 Upper Jurassic Upper Jurassic $-$-r gb-c-6-$ 204 AS-772 65.0 52.0 Callovian Callovian d-d-e gd-b-6-$ 204 AS-773 55.1 38.8 Tithonian Tithonian e-e-f gc-b-6-$ 204 AS-774 55,8 37.5 Portlandian Portlandian e-e-f b-b-6-S 204 AS-777 54.3 48.4 Tithonian Tithonian e-e-f gc-b-6-$ 204 AS-780 51.4 41.6 Tithonian Tithonian e-e-f gd-b-6-S 137; 204 AS-781 54.3 48.4 Tithonian Tithonian e-e-f gd-b-6-$ 204 AS-783 52.0 50.4 Tithonian Tithonian e-e-f gc-b-6-$ 204 AS-784 53.1 48.4 Tithonian Tithonian e-e-f gd-b-6-S 204 AS-785 53.0 72.0 Tithonian Tithonian e-c-f d-b-6-S 204 AS-786 54.3 48.4 Valanginian Valanginian f-f-g gd-c-6-$ 204 AS-787 53.2 48.5 Valanginian Valanginian f-f-g gc-c-6-$ 204 AS-788 54.3 48.4 Lower Crctaceous Lower Cretaceous $-$-g gd-c-6-S 204 AS-789 53.9 48.7 Lower Cretaccous Lower Cretaceous $-$-g gc-c-6-$ 204 AS-790 58.8 52.3 Valanginian Valanginian f-f-g gc-b-6-$ 204 AS-791 41.0 48.8 Lower Cretaceous Lower Cretaceous S-S-g gb-c-6-$ 204 AS-792 53.0 40.9 Lower Cretaceous Lower Cretaceous S-$-g gb-c-6-S 204 AS-793 52.5 48.0 Lower Cretaceous Lower Cretaceous S-$-g gd-c-6-$ 204 AS-794 53.8 43.2 Albian Albian S-h-g gc-b-3-S 204 AS-795 51.3 37.5 Albian Cenomanian S-h-g gb-c-6-5 204 AS-796 54.6 48.4 Banemian Barremian $-f-g g-c-6-S 204 AS-797 48.8 25.2 Albian Cenomanian S-h-g g-c-6-$ 204 AS-798 48.4 27.8 Cenomanian Cenomanian h-h-h gb-b-6-S 204 AS-799 48.4 27.8 Cenomanian Cenomanian h-h-h gb-b-6-S 204 AS-800 51.5 45.5 Cenomanian Cenomanian h-h-h gc-b-6-S 204 AS-801 51.4 45.8 Cenomanian Cenomanian h-h-h gc-b-6-$ 204 AS-802 51.3 48.2 Cenomanian Cenomanian h-h-h gc-b-6-$ 204 AS-803 51.5 45.2 Cenomanian Cenomanian h-h-h gc-b-6-S 204 AS.804 51.6 43.2 Cenomanian Cenomanian h-h-h gc-b-6-$ 204 AS-805 52.8 41.5 Cenomanian Cenomanian h-h-h gd-b-6-S 204 AS-806 51.2 143.1 Barremian Cenomanian S-$-g gd-d-6-S 204 AS-807 46.2 38.8 Cenomanian Cenomanian h-h-h gc-b-6-S 204 AS-808 39.9 67.7 Upper Cretaceous Upper Cretaceous $-$-h gc-c-6-S 132 AS-809 52.5 64.5 Upper Cretaceous Upper Cretaceous $-S-h b-c-6-S 236 AS-810 41.6 61.4 Cenomanian Campanian $-i-h gc-c-6-$ 139 AS-849 11.6 102.5 Lower Jurassic Lower Jurassic $-$-d gb-c-6-S 105 AS-887 16.6 106.2 Lower Jurassic Lower Jurassic S-S-d gb-c-6-$ 125 AS-888 40.5 -105.0 Albian Albian S-h-g gb-b-3-def 304 AS-895 45.9 86.5 Lower Cretaceous Lower Cretaceous S-S-g b-c-2-de 329 AS-9I 52.6 43.6 Santonian Santonian S-i-h ge-B-6-$ 219 AS-92 29.6 106.3 Lower Jurassic Lower Jurassic S-S-d gd-C-3-$ 88 AS-93 33.0 71.4 Upper Jurassic Lower Cretaceous S-S-f ge-D-3-$ 44 AS-94 23.7 69.0 Tithonian Tithonian e-e-f a-C-2-S 303; 24 AU-lOO -20.8 144.2 Aptian Aptian g-g-g gc-B-3-S 226; 196;296 AU-IOI -20.7 144.5 Aptian Aptian g-g-g ge-B-S-S 226; 296 AU-102 -20.7 143.1 Albian Albian S-h-g ge-B-S-S 226 AU-103 -28.1 136.8 Albian Albian S-h-g gc-B-S-S 226; 196 AU-I04 -26.6 148.2 Aptian Aptian g-g-g ge-B-S-S 226 AU-105 -20.9 143.9 Albian Albian S-h-g ge-B-S-S 226 AU-285 -42.6 -173.5 Upper Cretaceous Upper Cretaceous S-S-h g-C-S-S 229 AU-320 -31J 115.9 Upper Cretaceous Upper Cretaceous S-S-h a-c-3-f 159 AU-41 -20.0 1442 Albian Albian S-h-g d-$-3-S 305 AU-450 -422 173.3 Maastrichtian Maastrichtian j-j-h ge-b-3-de 310;306 AU-45I -43.2 172.8 Maastrichtian Maastrichtian j-j-h gd-b-3-de 310;306 AU-452 -42.6 173.5 Maastrichtian Maastrichtian j-j-h ge-b-3-de 310;306 305

TABLE G. 1 Marine Reptile Database Localities Age Range Lat Lon Maximum Minimum inicrv^ Reliability Reference

AU-453 -42.8 172.6 Maastrichtian Maastrichtian j-j-h gc-b-3-de 310; 306 AU-454 -43.1 172.6 Maastrichtian Maastrichtian j-j-h ge-b-3-de 310; 306 AU-455 -43.2 172.8 Maastrichtian Maastrichtian j-j-h gd-b-3-de 310;306 AU-456 -43.6 170.5 Maastrichtian Maastrichtian j-j-h gd-b-3-de 310; 306 AU-457 -42.8 173.3 Maastrichtian Maastrichtian j-j-h ge-b-3-de 310; 306 AU-509 -39.3 177.5 Campanian Maastrichtian $-i-h ge-c-3-S 3I5;3I6 AU-5I -20.0 144.2 Albian Albian $-h-g d-B-6-F 173;17 AU-635 -45.5 169.0 Upper Cretaceous Upper Cretaceous $-$-h gc-c-6-S 98 AU-68 -44.1 171.4 Camian Camian S-a-c g*-B-4-S 55; 97 AU-69 -46.1 168.5 Camian Camian $-a-c g'-A-2-S 55 AU-70 -40.9 176.1 Albian Albian $-h-g g*-A-2-$ 97 AU-707 -21.7 165.7 Norian Norian b-b-c gb-a-2-f 54 AU-740 -46.4 169.8 Camian Camian S-a-c gb-b-6-S 48; 99 AU-753 -45.8 168.1 Anisian Anisian S-a-b gc-b-6-S 48 AU-825 -35.9 173.9 Ccnomanian Cenomanian h-h-h gb-c-6-$ 99 AU-826 -45.8 168.1 Middle Triassic Middle Triassic $-$-b gc-c-6-$ 99 AU-827 -43.3 171.9 Maastrichtian Maastrichtian j-j-h gb-b-3-S 99 AU-828 -23.6 150.4 Lower Jurassic Lower Jurassic $-$-d gb-c-3-S 196 AU-829 -12.3 130.9 Albian Albian $-h-g a-b-l-f 201 AU-84 -25.7 150.4 Lower Jurassic Lower Jurassic S-S-d c-C-3-$ 295; 196 AU-856 -29,0 134.7 Aptian Aptian g-g-g gb-b-3-$ 197 AU-857 -30.5 137.2 Aptian Aptian g-g-g gb-b-3-$ 197 AU-97 -30.8 143.1 Aptian Aptian g-g-g ge-B-S-S 226 AU-98 -21.5 142.9 Aptian Aptian g-g-g ge-B-S-S 226 AU-99 -20.9 143.6 Albian Albian $-h-g g'-B-S-S 226 EU-106 49.4 11.9 Aalcnian Aalenian S-d-e ge-A-S-S 312 EU-I07 50.9 -0.3 Ccnomanian Cenomanian h-h-h gc-C-S-S 229;34 EU-108 51.2 0.8 Turonian Campanian S-i-h gc-C-S-S 229 EU-109 51.1 1.2 Albian Albian $-h-g ge-B-S-S 229;34 EL'-110 52.2 0.1 Albian Albian $-h-g ge-B-S-S 229;34 EU-111 51.3 0.5 Aptian Aptian g-g-g ge-B-S-S 229;34 EU-112 50.8 0.6 Valanginian Valanginian f-f-g ge-B-S-S 229;34 EU-113 52.3 0.3 Lower Cretaceous Lower Cretaceous S-$-g g-C-S-S 229 EU-I14 50.8 0.2 Bcrriosion Barrcmian S-f-g gc-B-S-S 229;34 EU-115 50.5 -2.4 Portlandian Portlandian e-e-f ge-B-S-S 229;34 EU-n6 51.9 -0.9 Portlandian Portlandian e-e-f gc-B-S-S 229 EU-n7 50.6 -2.6 Kimmeridgian Kimmeridgian $-e-f ge-B-S-S 229;34 EU-118 50.6 -2.1 Kimmeridgian Kimmeridgian S-c-f ge-B-S-S 229;34 EU-119 52.4 -0.3 Kimmeridgian Kimmeridgian S-e-f gcd-B-S-S 229;34 EU-12 50.0 9.5 Anisian Anisian S-a-b b-A-2-F 254 EU-120 51.3 -1.9 Kimmeridgian Kimmeridgian S-e-f gd-B-S-S 229;34 EU-12I 51.8 -1.2 Kimmeridgian Kimmeridgian S-e-f gd-B-S-S 229;34 EU-I22 51.7 -1.2 Kimmeridgian Kimmeridgian S-e-f g-B-S-S 81; 229; 34 EU-123 52.6 -0.2 Callovian Oxfordian S-d-e ge-B-S-S 229;34 EU-124 52.1 -0.4 Callovian Oxfordian S-d-e ge-B-S-S 229;34 EU-125 51.8 -1.2 Callovian Oxfordian S-d-e ge-B-S-S 229;34 EU-126 50.6 -2.4 Callovian Oxfordian S-d-e ge-B-S-S 229;34 EU-127 51.8 -1.3 Callovian Oxfordian S-d-e gb-B-S-S 81; 229; 34 EU-I28 56.9 -6.2 Bathonian Bathonian S-d-e ge-B-S-S 229;34 EU-129 53.4 -0.3 Kimmeridgian Kimmeridgian S-e-f ge-B-S-S 81; 229; 34 EU-130 51.5 -1.8 Kimmeridgian Kimmeridgian S-e-f ge-B-S-S 229; 34 EU-131 51.8 -1.2 Kimmeridgian Kimmeridgian S-c-f g-B-S-S 229; 34 EU-132 54.5 -0.6 Toarcian Toarcian S-c-d g-A-3-S 229;35 EU-133 54.6 -0.9 Toarcian Toarcian S-c-d g-A-3-S 229;35 EU-134 50.7 -2.9 Sinemurian Sinemurian S-c-d ge-A-S-S 229 EU-135 51.5 -2.8 Sinemurian Sinemurian S-c-d gc-A-S-S 229 EU-136 50.7 -2.9 Hettangian Hettangian S-c-d ge-A-S-S 229 EU-137 51.6 -2.4 Rhaetian Lias S-b-c e-D-4-S 229; 284 EU-138 54.5 -0.7 Toarcian Toarcian S-c-d ge-A-3-S 229;35 306

TABLE G.l Marine Reptile Database Localities Age Range Lat Lon Maximum Minimum Interim Reliability Reference

EU-139 52.2 -0.9 Lower Jurassic Lower Jurassic $-$-d ge-C-S-S 229 EU-140 54.5 -0.6 Toarcian Toarcian S-c-d g-A-3-$ 229;35 EU-141 54.5 -0.6 Toarcian Toarcian S-c-d g-A-3-S 229;35 EU-142 52.9 -0.9 Sinemurian Sinemurian $-c-d ge-A-$-S 229 EU-\43 51.4 -2.4 Sinemurian Sinemurian $-c-d geb-A-S-$ 229 EU-144 50.7 -2.9 Sinemurian Sinemurian S-c-d ge-A-$-$ 229 EU-145 51.1 -2.8 Hettangian Hettangian $-c-d gec-B-S-S 81; 229 EU-146 52.2 -1.4 Hettangian Hettangian S-c-d ge-A-S-S 229 EU-147 51.4 -2.4 Hettangian Hettangian S-c-d gec-C-S-S 81; 229 EU-148 51.6 -2.6 Rhaetian Rhaetian $-b-c ge-B-3-df 229;284;34 EU-149 51.8 -1.4 Bathonian Bathonian S-d-e ge-B-S-S 229;34 EU-150 50.5 3.8 Upper Cretaceous Upper Cretaceous $-$-h gec-C-$-$ 229 EU-151 50.4 3.9 Upper Cretaceous Upper Cretaceous S-S-h ge-C-$-$ 229 EU-152 50.6 4.7 Upper Cretaceous Upper Cretaceous S-$-h gec-C-$-S 229 EU-153 49.3 4.9 Albian Albian $-h-g ge-C-S-S 229 EU-154 49.4 5.4 Albian Albian S-h-g geb-C-$-S 229 EU-155 52.2 7.1 Neocomian Neocomian S-f-g c-C-$-S 229 EU-156 52.3 9.0 Neocomian Neocomian $-f-g e-C-S-S 229 EU-157 52.0 8.4 Neocomian Neocomian S-f-g gec-C-S-S 229 EU-159 50.7 1.6 Portlandian Portlandian e-e-f ge-A-S-S 229 EU-160 50.8 1.6 Portlandian Portlandian e-e-f gb-A-S-S 229 EU-I6I 50.7 1.6 Kimmeridgian Kimmeridgian S-e-f ge-A-S-S 229 EU-162 50.7 1.6 Kimmeridgian Kimmeridgian S-e-f ge-A-S-S 229 EU-164 49.1 -0.2 Callovian Callovian d-d-e ge-A-2-df 229; 103 EU-165 50.7 1.6 Bathonian Bathonian $-d-c gc-B-$-S 229 EU-167 49.5 0.1 Kimmeridgian Kimmeridgian $-e-f e-B-S-S 229 EU-168 50.7 1.6 Oxfordian Oxfordian S-e-f d-B-S-S 229 EU-i69 46.9 2.8 Callovian Callovian d-d-e gec-B-S-S 229 EU-170 48.7 -0.1 Bathonian Bathonian S-d-e ge-B-S-S 229 EU-I7I 48.8 -6.2 Pliensbachian Pliensbachian c-c-d ge-A-S-S 229 EU-172 50.0 5.5 Sinemurian Sinemurian S-c-d ge-a-3-df 229;117 EU-173 49.4 -0.6 Lower Jurassic Lower Jurassic S-S-d gc-C-S-S 229 EU-174 47.0 4.3 Rhaetian Rhaetian S-b-c gc-B-S-S 229 EU-175 48.6 6.5 Middle Triassic Middle Triassic S-S-b gc-C-S-S 229:21 EU-176 38.7 -9.2 Cenomanian Cenomanian h-h-h ge-B-S-S 229 EU-177 40.2 -8.8 Pliensbachian Toarcian S-c-d ge-C-S-S 229 EU-178 50.1 14.5 Turonian Turonian S-i-h g»-B-$-S 229 EU-179 50.7 23.2 Turonian Turonian S-i-h gd-B-S-S 229 EU-180 50.5 14.8 Turonian Turonian S-i-h gc-B-3-S 309;229 EU-181 49.8 14.4 Turonian Turonian S-i-h gc-B-3-S 309;229 EU-182 50.6 13.8 Turonian Turonian S-i-h gec-B-3-$ 309:229 EU-183 50.0 16.5 Turonian Turonian S-i-h gb-B-S-S 229 EU-184 52.4 9.6 Lower Cretaceous Lower Cretaceous S-$-g ge-C-S-S 229 EU-185 46.8 6.5 Aptian? Aptian? g-g-g ge-C-S-S 229 EU-186 51.9 10.3 Lower Cretaceous Lower Cretaceous S-S-g ge-C-S-S 229 EU-187 52.1 10.2 Lower Cretaceous Lower Cretaceous S-S-g ge-C-S-S 229 EU-189 47.3 8.2 Oxfordian Oxfordian s-e-f gec-B-S-S 229 EU-190 48.1 8.6 Callovian Callovian d-d-e gec-A-S-S 229 EU-191 49.1 9.6 Oxfordian? Oxfordian? $-e-f ge-C-S-S 229 EU-192 51.4 22.8 Portlandian Portlandian e-e-f ge-B-S-S 229 EU-193 48.6 9.5 Toarcian Toarcian S-c-d ge-C-S-S 229 EU-195 49.2 8.6 Sinemurian Sinemurian S-c-d gc-A-S-S 229 EU-I96 48.7 9.1 Sinemurian Sinemurian s-c-d ge-A-S-S 229 EU-197 48.6 9.0 Sinemurian Sinemurian S-c-d ge-A-S-S 229 EU-198 51.9 11.1 Upper Triassic Upper Triassic S-S-c c-C-S-S 229 EU-199 50.0 11.6 Middle Triassic Middle Triassic S-S-b ge-C-S-S 229 EU-20 49.5 9.4 Anisian Anisian S-a-b ge-A-2-F 241 EU-200 51.9 11.1 Hettangian Hettangian S-c-d d-A-S-S 229 EU-201 49.1 9.6 Rhaetian Rhaetian S-b-c ge-B-S-S 229 307

TABLE G.l Marine Reptile Database Localities Age Range Lat Lon Maximum Minimum Int^^ Reliability Reference

EU-203 48.5 9.2 Toarcian Toarcian $-c-d ge-C-S-S 229 EU-204 48.1 8.7 Lower Jurassic Lower Jurassic $-$-d ge-C-S-S 229 EU-205 48.9 10.1 Sinemurian Sinemurian $-c-d ge-C-S-S 229 EU-206 54.0 19.2 Campanian Campanian $-i-h ge-b-S-S 229; 22 EU-207 56.1 13.9 Campanian Campanian $-i-h gc-A-S-S 229 EU.21 51.5 12.0 Anisian Anisian $-a-b gb-B-2-F 241 EU-2I0 54.0 19.0 Turonian Campanian $-i-h gb-b-S-S 229:22 EU-23 50.5 18.0 Anisian Anisian S-a-b g*-B-2-F 241 EU-244 78.4 16.5 Oxfordian? Oxfordian? $-e-f gc*-C-$-$ 229 EU-245 78.3 15.8 Upper? Jurassic Upper? Jurassic $-$-f gb-D-S-S 229 EU-246 78.4 15.9 Portlandian? Portlandian? e-e-f gb-C-S-S 229 EU-248 78.3 15.8 Ladinian Ladinian $-a-b gb-b-2-ef 229; 72 EU-25 50.4 -3.2 Sinemurian Sinemurian $-c-d b-A-2-$ 34; 285 EU-298 44.9 9.3 Santonian Campanian $-i-h ge-b-3-S 239 EU-299 48.6 9.5 Toarcian Toarcian $-c-d gc-b-2-S 177 EU-300 48.6 9.5 Toarcian Toarcian $-c-d ge-b-2-$ 177 EU-30I 48.6 9.5 Toarcian Toarcian $-c-d ge-b-2-S 177 EU-302 48.6 9.5 Toarcian Toarcian $-c-d ge-b-2-S 177 EU-303 48.6 9.5 Toarcian Toarcian $-c-d ge-b-2-S 177 EU-304 48.6 9.5 Toarcian Toarcian $-c-d ge-b-2-$ 177 EU-305 48.6 9.5 Toarcian Toarcian S-c-d ge-b-2-S 177 EU-306 48.6 9.5 Toarcian Toarcian $-c-d ge-b-2-$ 177 EU-307 48.6 9.5 Toarcian Toarcian $-c-d ge-b-2-S 177 EU-308 48.6 9.5 Toarcian Toarcian $-c-d ge-b-2-$ 177 EU-309 48.6 9.5 Toarcian Toarcian S-c-d ge-b-2-S 177 EU-3I0 49.1 9.6 Toarcian Toarcian $-c-d gd-b-2-S 177 EU-3II 48.7 9.7 Toarcian Toarcian $-c-d ge-b-2-$ 177 EU-312 48.1 8.7 Toarcian Toarcian S-c-d d-b-2-S 177 EU-36 52.5 •0.3 Callovian Callovian d-d-e c-A-l-DF 160 EU-395 48.9 11.9 Cenomanian Cenomanian h-h-h ge-a-l-df 29 EU-397 43.2 23.7 Maastrichtian Maastrichtian j-j-h ge-b-2-f 212 EU-417 50.5 -2.4 Kimmeridgian Kimmeridgian $-e-f gb-a-2-S 41:221 EU-418 38,7 -7.1 Sinemurian Sinemurian S-c-d ge-b-6-S 327 EU-»20 40.2 -8.8 Pliensbachian Toarcian? S-c-d ge-c-6-S 327 EU-421 40,4 -8.5 Pliensbachian Toarcian? S-c-d gd-c-6-S 327 EU-422 40.3 -8.6 Pliensbachian Pliensbachian c-c-d gd-c-6-S 327 EU-423 40.4 -8.5 Pliensbachian? Pliensbachian? c-c-d ge-c-6-S 327 EU-424 39.8 -9.0 Pliensbachian? Pliensbachian? c-c-d gc-c-6-$ 76 EU-425 40.1 -8.3 Toarcian Toarcian S-c-d ge-a-4-S 76 EU-426 39.6 -8.4 Aalenian Aalenian S-d-e ge-b-6-S 76 EU-427 40.3 -8.2 Pliensbachian Pliensbachian c-c-d ge-b-6-$ 76 EU-428 47.7 6.3 Toarcian Toarcian S-c-d ge-a-4-de 70 EU-429 47.7 6.3 Toarcian Toarcian S-c-d ge-a-4-de 70 EU-430 47.5 6.4 Toarcian Toarcian S-c-d ge-a-4-de 70 EU-431 47.2 5.9 Toarcian Toarcian S-c-d ge-a-4-de 70 EU-432 47.4 6.4 Toarcian Toarcian S-c-d ge-a-4-de 70 EU-433 47.0 5.9 Toarcian Toarcian S-c-d ge-a-4-de 70 EU-434 46.4 5.8 Toarcian Toarcian S-c-d ge-a-4-de 70 EU-435 50.7 -2.9 Lower Jurassic Lower Jurassic $-S-d gb-c-6-S 84 EU-436 50.9 -2.6 Bathonian Callovian S-d-e ge-b-3-S 84 EU-437 50.6 -2.1 Kimmeridgian Kimmeridgian S-e-f gc-b-3-S 84 EU-438 50.6 -2.4 Oxfordian Oxfordian $-e-f ge-b-3-S 85 EU-439 50.6 -2.0 Kimmeridgian Kimmeridgian S-e-f ge-b-3-$ 292 EU-440 50.6 -2.0 Kimmeridgian Kimmeridgian $-e-f ge-a-3-S 292 EU-441 50.6 -2.0 Kimmeridgian Kimmeridgian S-e-f ge-a-3-S 292 EU-442 50.6 -2.0 Kimmeridgian Kimmeridgian S-e-f ge-b-3-S 292 EU-443 50.6 -2.0 Kimmeridgian Kimmeridgian S-e-f ge-a-3-S 292 EU-444 50.6 -2.0 Kimmeridgian Kimmeridgian S-e-f ge-a-3-S 292 EU-445 50.6 -2.0 Kimmeridgian Kimmeridgian S-e-f ge-a-3-S 292 308

TABLE G.l Marine Reptile Database Localities Age Range LoC Lat Lon Maximum Minimum \t3D Reliability Rererencc

EU-446 50.6 -2.0 Kimmeridgian Kimmeridgian $-e-f ge-b-3-$ 292 EU-448 50.8 1.7 Oxfordian Oxfordian S-e-f ge-a-6-$ 32 EU-449 50.6 -2.0 Upper Jurassic Lower Crctaceous $-$-f ge-d-6-S 83 EU-458 50.8 5.7 Maastrichtian Maasuichtian j-j-h gc-b-6-S 237 EU-459 50.8 5.7 Maastrichtian Maastrichtian j-j-h ge-b-6-S 237 EU-46 52.2 0.2 Cenomanian Cenomanian h-h-h e-B-3-F 17 EU-460 50.8 5.6 Maastrichtian Maastrichtian j-j-h ge-b-6-S 237 EU-461 50.8 5.6 Maastrichtian Maastrichtian j-j-h ge-b-6-$ 237 EU-462 50.4 3.9 Maastrichtian Maastrichtian J-j-h ge-b-6-S 237 EU-463 50.4 4.0 Maastrichtian Maastrichtian j-j-h ge-b-6-$ 237 EU-464 50.5 3.8 Maastrichtian Maastrichtian j-j-h ge-b-6-$ 237 EU-465 50.4 3.9 Maastrichtian Maastrichtian j-j-h ge-b-6-$ 237 EU-466 50.8 5.7 Maastrichtian Maastrichtian j-j-h gb-b-6-$ 237 EU-47 52.4 9.8 Aptian Aptian g-g-g e»-B-4-F 17 EU-475 50.6 •2.4 Kimmeridgian Kimmeridgian $<-f ge-b-3-S 80 EU-479 50.7 -2.7 Lower Jurassic Lower Jurassic $-$-<1 ge-c-6-$ 81 EU-48 52.4 9.8 Aptian Aptian g-g-g e*-C-6-F 17 EU-480 50.5 -2.5 Kimmeridgian Kimmeridgian $-€-f ge-b-3-$ 81 EU-481 50.6 -2.4 Portland ian Portlandian e-e-f gc-b-3-S 81 EU-482 50.6 -2.4 Portlandian Portlandian e-e-f gc-b-3-S 81 EU-483 51.6 -1.6 Portlandian Portlandian e-e-f ge-b-3-$ 81 EU-484 50.6 -2.5 Callovian Oxfordian S-d-e ge-b-3-$ 81 EU-485 51.2 -0.8 Callovian Oxfordian $-d-« ge-b-3-S 81 EU-486 50.6 -2.4 Kimmeridgian Kimmeridgian $-«-f ge-b-3-S 81 EU-487 50.7 -1.5 Kimmeridgian Kimmeridgian $-e-f ge-b-3-S 81 EU-488 50.6 -2.4 Portlandian Portlandian e-«-f gc-b-3-$ 81 EU-489 51.0 -2.2 Kimmeridgian Kimmeridgian S-e-f ge-b-3-$ 81 EU-490 50.8 -2.4 Kimmeridgian Kimmeridgian S-e-f gc-b-3-S 81 EU-491 69.2 16.1 Oxrordian Kimmeridgian S-e-f ge-b-3-f 218 EU-507 51.2 -3.2 Sinemurian Sinemurian S-c-d ge-a-4-$ 178 EU-5I2 51.8 -1.2 Callovian Oxfordian S-d-e gc-b-3-S 82 EU-513 50.8 -2.4 Kimmeridgian Kimmeridgian S-e-f gc-b-3-S 82 EU-514 51.3 -1.9 Kimmeridgian Kimmeridgian S-c-f gc-b-3-S 82 EU-515 50.7 -2.9 Albian Albian S-h-g gb-b-3-S 82 EU-516 51.0 -2.3 Kimmeridgian Kimmeridgian S-c-f gb-b-3-$ 82 EU-527 47.0 6.9 Hauterivian Hauterivian S-f-g e-c-6-$ 309 EU-528 50.1 14.3 Turonian Turonian S-i-h ge-c-3-$ 309 EU-53 50.9 1.9 Cenomanian Cenomanian h-h-h b*-A-l-DEF 17 EU-530 50.8 0.0 Turonian Campanian S-i-h ge-c-6-S 309 EU-54 45.0 4.7 Toarcian Toarcian S-c-d c»-A-I-DEF 91 EU-541 47.8 13.3 Sinemurian Pliensbachian S-c-d gc-c-6-d 134:234 EU-542 46.6 11.7 Ladinian Ladinian S-a-b gb-b-3-S 142 EU-543 48.0 8.2 Callovian Callovian d-d-c gd-b-3-S 263 EU-55 48.6 9.5 Toarcian Toarcian S-c-d gb-A-l-S 129 EU-553 52.7 1.0 Turonian Campanian S-i-h gc-c-3-f 27 EU-554 42.9 2.4 Santonian Santonian S-i-h gb-b-3-f 27 EU-555 42.9 2.4 Santonian Santonian S-i-h gb-b-3-f 27 EU-556 54.0 19.1 Santonian Santonian S-i-h d-b^-f 27 EU-557 49.9 2.8 Santonian Santonian S-i-h gb-b-3-f 27 EU-558 50.4 4.9 Santonian Santonian S-i-h d-b-3-f 27 EU-559 49.3 4.0 Campanian Campanian S-i-h gb-b-3-f 27 EU-56 48.6 9.5 Toarcian Toarcian S-c-d gb-A-I-S 129 EU-560 48.3 3.2 Campanian Campanian S-i-h gb-b-3-f 27 EU-561 44.2 4.5 Campanian Campanian S-i-h gd-b-3-f 27 EU-562 43.4 -6.2 Campanian Campanian S-i-h gb-b-3-f 27 EU-563 45.5 -0.8 Campanian Campanian S-i-h gb-b-3-f 27 EU-564 54.0 19.1 Campanian Campanian S-i-h d-b-4-f 27 EU-565 56.2 15.5 Campanian Campanian S-i-h gb-b-4-f 27 EU-566 48.8 2.2 Campanian Campanian S-i-h d-b-3-f 27 309

TABLE G. I Marine Reptile Database Localities Age Ranee Lat Lon Maximum Minimum Interval Reliability Reference

EU-567 51.4 7.5 Campanian Campanian $-i-h d-b-4-f 27 EU-568 50.5 3.9 Campanian Campanian S-i-h d-b-3-r 27 EU-569 54.2 -6.3 Campanian Maastrichtian S-i-h gc-b-3-f 27 EU-57 48.6 9.5 Toarcian Toarcian S-c-d gb-A-l-S 129 EU-S70 50.5 3.9 Maastrichtian Maastrichtian j-j-h d-b-3-f 153:27 EU-571 50.5 3.9 Maastrichtian Maastrichtian j-j-h d-b-3-r 22; 27 EU-572 50.9 5.6 Maastrichtian Maastrichtian j-j-h d-b-3-r 200; 27) EU-573 43.3 •0.6 Maastrichtian Maastrichtian j-j-h gb-b-3-r 27 EU-574 42.7 -2.6 Maastrichtian Maastrichtian j-j-h gb-b-6-f 27 EU-575 55.3 12.4 Maastrichtian Maastrichtian j-j-h gb-b-6-f 27 EU-576 50.5 3.9 Maastrichtian Maastrichtian j-j-h d-b-3-f 27 EU-577 50.5 3.9 Campanian Campanian $-i-h d-b-3-f 22; 27 EU-58 48.2 8.6 Toarcian Toarcian $-c-d e*-A-l-$ 129 EU-582 44.3 6.3 Aalenian Aalenian S-d-c gc-a-2-S 7 EU-584 48.4 3.0 Cenomanian Cenomanian h-h-h gc-b-6-S 19 EU-59 48.6 9.5 Toarcian Toarcian S-c-d gb-A-l-S 129 EU-60 43.5 1.4 Albian Albian S-ii-g g*-C-7-F 23 EU-609 52.7 -1.2 Rhaetian Hettangian $-b-c gb-c-7-S 74 EU-6I 49.7 -5.8 Sinemurian Sinemurian $-c-d g*-A-2-DF 115 EU-610 45.6 10.4 Maastrichtian Maastrichtian J-j-h gb-b-3-S 276 EU-6II 78.3 16.8 Spathian Spathian $-a-a gb-b-4-S 72; 166 EU-612 77.9 23.5 Camian Camian $-a-c b-c-2-cf 72 EU-613 76.7 25.5 Rhaetian Rhaetian $-b-c b-b-3-cf 72; 280 EU-614 77.5 22.0 Ladinian Ladinian S-a-b b-b-2-cf 72 EU-6I5 77.7 21.5 Ladinian Ladinian $-a-b b-b-2-ef 72 EU-616 77.5 22.5 Ladinian Ladinian S-a-b b-b-2-ef 72 EU-617 78.1 23.0 Ladinian Ladinian $-a-b b-b-2-ef 72 EU-618 78.4 21.9 Ladinian Ladinian $-a-b b-b-2-ef 72 EU-619 78.3 16.3 Anisian Ladinian S-a-b b-c-3-ef 72 EU-620 78.1 17.8 Anisian Ladinian S-a-b b-c-3-cf 72 EU-621 78.2 17.3 Anisian Ladinian S-a-b b-c-3-cr 72 EU-622 78.6 14.1 Anisian Ladinian S-a-b b-c-3-ef 72 EU-623 78.4 15.3 Smithian Spathian S-a-a b-b-3-cf 72 EU-624 78.6 14.1 Smithian Spathian S-a-a b-b-3-ef 72 EU-625 78.2 18.0 Smithian Spathian S-a-a b-b-3-cf 72 EU-626 77.9 23.5 Smithian Spathian S-a-a b-b-3-ef 72 EU-627 78.4 21.9 Smithian Spathian S-a-a b-b-3-cf 72 EU-631 43.8 3.8 Toarcian Toarcian S-c-d gb-b-l-d 143 EU-632 49.2 6.0 Bajocian Bajocian S-d-e gb-a-3-r 116 EU-633 49.8 16.3 Turonian Turonian S-i-h gb-b-6-S 326 EU-634 50.5 3.8 Maastrichtian Maastrichtian j-j-h gc-b-3-S 147 EU-637 43.1 5.9 Rhaetian Rhaetian S-b-c gb-b-6-f 21 EU-639 47.2 5.5 Rhaetian Rhaetian S-b-c gb-b-6-r 21 EU-640 48.0 5.5 Rhaetian Rhaetian S-b-c gb-b-6-f 21 EU-641 41.3 1.2 Ladinian Ladinian S-a-b gb-b-3-f 261 EU-654 78.9 28.2 Rhaetian Rhaetian S-b-c gb-b-2-df 321 EU-66 45.9 9.0 Anisian Ladinian S-a-b gb-B-3-F 141;255 EU-663 49.5 9.0 Anisian? Anisian? S-a-b gc-b-2-df 121 EU-665 49.5 0.1 Kimmeridgian Kimmeridgian S-e-f gb-b-6-$ 25 EU-666 43.8 3.2 Toarcian Toarcian S-c-d gb-a-4-S 258 EU-667 49.4 0.0 Callovian Callovian d-d-c gb-a-3-$ 26 EU-668 78.4 15.3 Ladinian Ladinian S-a-b d-a-2-f 169 EU-669 47.5 8.3 Upper Lias Upper Lias S-c-d gb-c-6-S 318 EU-674 57.6 -6.2 Callovian Callovian d-d-e gb-b-3-d 69 EU-679 56.4 12.9 Campanian Campanian S-i-h gb-b-6-S 225 EU-681 44.0 3.0 Toarcian Toarcian S-c-d gb-a-4-$ 267 EU-689 44.2 5.8 Valanginian Valanginian r-f-g gb-a-4-$ 294 EU-690 44.2 5.7 Valanginian Valanginian f-f-g gb-a-4-$ 294 EU-703 48.9 11.0 Tithonian Tithonian e-e-f gb-c-3-de 30 310

TABLE G.l Marine Reptile Database Localities Age Range Lat Lot! Maximum Minimum interim Reliability Reference

EU-704 50.9 21.8 Maastrichtian Maastrichtian j-j-h gc-a-4-f 289 EU-705 51.4 22.0 Maastrichtian Maastrichtian j-j-h gc-a-4-f 289 EU-713 55 0 -5.9 Lower Lias Lower Lias $-c-d gb-c-4-S 133 EU-72 52.6 0.4 Kimmeridgian Kimmeridgian J-e-f g*-A-2-$ 176 EU-720 51.1 -2.3 Oxfordian Oxfordian $-e-f b-a-3-f 302 EU-733 53.6 11.5 Lower Jurassic Lower Jurassic S-S-d g-c-6-f 199 EU-735 46.7 9.8 Norian Rhaetian $-b-c gc-c-3-S 48 EU-742 46.8 13.8 Camian Camian S-a-c gc-b-3-$ 48 EU-743 45.0 28.9 Middle Triassic Middle Triassic $-$-b gb-c-6-S 48 EU-744 50.2 18.8 Ladinian Ladinian $-a-b gc-b-2-$ 48 EU-745 49.2 9.3 Ladinian Ladinian $-a-b ed-b-2-S 48 EU-746 49.2 7.0 Ladinian Ladinian $-a-b gc-b-2-S 48 EU-749 47.2 15.2 Anisian Ladinian S-a-b gc-c-3-S 48 EU-750 45.9 8.9 Anisian Ladinian $-a-b gb<-3-S 232;48 EU-751 47.3 8.2 Anisian Anisian $-a-b gc-b-3-S 48 EU-752 48.0 8.2 Anisian Anisian S-a-b gd-b-2-S 48 EU-759 78.2 17.0 Ladinian Ladinian $-a-b d-b-2-f 256 EU-76 43.6 10.4 Cenomanian Cenomanian h-h-h g«-c-2-S 243; 19 EU-79 49.2 0.0 Albian Albian $-h-g ge-A-4-S 79 EU-80 49.6 -1.6 Kimmeridgian Kimmeridgian $-e-f gc-B-6-S 78 EU-8I 39.8 -9.0 Pliensbachian Pliensbachian c-c-d ge-B-4-$ 76 EU-811 43.8 5,6 Hautcrivian Hauterivian $-f-g gb-b-6-$ 100 EU-812 49.6 5.8 Pliensbachian Pliensbachian c-c-d gb-b-3-$ 114 EU-814 45.6 10.2 Anisian Ladinian S-a-b gb-c-3-S 38 EU-8I5 43.4 12.9 Kimmeridgian Tithonian S-e-f gb-c-3-f 92 EU-85 52.5 16.0 Oxrordian Oxfordian $-e-f C-B-6-S 163 EU-86 50.8 19.1 Oxrordian Oxfordian S-e-f gb-B-3-S 163 EU-87 56.0 14.0 Campanian Campanian S-i-h g-B-6-S 228 EU-88 59.2 14.1 Campanian Campanian $-i-h ge-B-6-$ 228 EU-89 56.2 15.5 Campanian Campanian $-i-h g*-B-6-$ 225; 228: EU-90 61.3 13.8 Upper Cretaceous Upper Cretaceous S-$-h g-D-6-S 228 EU-95 56.1 14.5 Campanian Campanian S-i-h ge-B-4-S 230 EU-96 78.0 18.2 Callovian Oxfordian S-d-e gc-C-3-EF 113 NA-I 38.9 -117.6 Camian Camian S-a-c b-A-2-DEF 273; 275: NA-10 69.0 -129.0 Campanian Campanian S-i-h C-B-3-DE 233 NA-11 69.0 -129.0 Maastrichtian Maastrichtian j-j-h C-B-2-DE 233 NA-I 3 56.0 -123.0 Camian Norian S-a-c b-A-3-F 188 NA-14 56.0 -123.0 Norian Norian b-b-c b-A-I-F 188 NA-I 5 56.0 -123.0 Norian Norian b-b-c b-C-l-F 183;188 NA-I 6 56.0 -123,0 Upper Triassic Upper Triassic S-S-c b-C-3-F 188 NA-17 56.0 -123.0 Norian Norian b-b-c b-C-3-F 180; 188 NA-I 8 56.0 -123,0 Norian Norian b-b-c b-a-3-F 185; 188 NA-243 70.7 -26,8 Kimmeridgian Kimmeridgian S-e-f gb-B-S-S 229 NA-249 50.8 -111,5 Maastrichtian Maastrichtian j-j-h gc-B-S-S 229 NA-250 43.5 -99,4 Campanian Campanian S-i-h gb-A-S-S 229 NA-25I 38.9 -101,6 Campanian Campanian S-i-h gb-A-S-S 229 NA-252 42.5 -97.0 Coniacian Campanian S-i-h d-C-S-S 229 NA-2S3 35.8 -101.8 Coniacian Campanian S-i-h d»-C-S-S 229 NA-254 43.3 -103.8 Coniacian Campanian S-i-h e-C-S-S 307; 229 NA-255 38.9 -I0I.6 Campanian Campanian S-i-h gb-b-S-S 229; 56 NA-256 39.5 -97.6 Coniacian Coniacian i-i-h d*-b-$-$ 309:229 NA-257 32.6 -97.0 Turonian Turonian S-i-h d»-B-S-S 229 NA-258 37.4 -102.8 Cenomanian Turonian S-h-h gb-B-S-S 307; 229 NA-259 45.0 -104.4 Cenomanian Turonian S-h-h gb-C-S-S 229 NA-260 34.0 -93^ Upper? Upper? Cretaceous S-S-h d»-D-S-S 229 Cretaceous NA-261 38.9 -101.6 Campanian Campanian S-i-h gb-B-S-S 229; 56 NA-262 38.9 -97J Turonian Turonian S-i-h d»-B-S-S 229:56 NA-263 39.6 -97.7 Turonian Turonian S-i-h gb-B-S-S 229:56 311

TABLE G.l Marine Reptile Database Localities Age Range IP Lat Lot! Maximum Minimum Reliability Reference

NA-264 39.3 -97.8 Cenomanlan Turonian $-h-h c-B-S-S 229 NA-265 46.6 -98.7 Turonian Turonian $-i-h gb-B-S-S 229; 56 NA-266 44.2 -99.6 Campanian Campanian $-i-h d-B-S-S 229 NA-267 50.2 -96.9 Upper Cretaceous Upper Cretaceous S-$-h gb-C-S-S 307;229 NA-268 39.0 -100.5 Coniacian Coniacian i-i-h d»-B-S-$ 229 NA-269 46.0 -IIO.I Maastrichtian Maastrichtian j-j-h gd-B-S-S 307; 229 NA-270 48.0 -105.0 Campanian Campanian $-i-h d-A-S-S 229 NA-271 37.6 -99.3 Albian Albian $-h-g d*-B-S-S 307; 229 NA-272 37,2 -99.8 Aptian Albian 5-g-g c-B-S-S 229 NA-273 41.8 -105.7 Oxfordian Oxfordian $-d-e d*-B-S-S 229 NA-274 42.2 -106.4 Oxrordian Oxfordian $-d-e gc-B-S-S 111; 229 NA-275 43.0 -106.7 Kimmeridgian Portlandian $-d-c d*-B-S-S 229 NA-276 39.8 -75.4 Maastrichtian Maastrichtian j-j-h e-B-S-S 307; 229 NA-277 17,3 -97.7 Lower Cretaceous Lower Crctaceous S-$-g gb-C-S-S 307;229 NA-278 36.7 -120.7 Campanian Maastrichtian S-i-h gb-B-S-S 229 NA-279 35.3 -120.7 Portlandian Portlandian e-e-f d*-B-S-S 229 NA-286 69.3 -128.2 Upper Crctaceous Upper Cretaceous $-$-h a-A-3-DF 247 NA-287 69.3 -128.2 Maastrichtian Maastrichtian j-j-h a-B-3-DF 247 NA-288 49.2 -98.1 Campanian Campanian S-i-h gb-A-2-S 206 NA-289 54.5 -120.7 Griesbachian Spathian S-a-a a-C-2-S 209 NA-290 49.7 -125.1 Santonian Campanian S-i-h a-A-2-S 208 NA-297 53.6 -113.5 Maastrichtian Maastrichtian j-j-h e*-c-S-S 253 NA-318 40.0 -74.8 Maastrichtian Maastrichtian j-j-h gb-a-3-S 13 NA-319 36.9 -107.0 Campanian Campanian S-i-h gc-c-3-r 157 NA-321 38.9 -99.3 Turonian Turonian S-i-h gc-b-3-S 162 NA-322 38.9 -99.3 Turonian Turonian S-i-h gc-b-3-S 162 NA-323 39.3 -98.9 Turonian Turonian S-i-h gb-b-3-S 162 NA-324 39.7 -75.2 Maastrichtian Maastrichtian j-j-h gb-b-2-S 223 NA-325 40.3 -74.2 Maastrichtian Maastrichtian j-j-h gb-b-2-S 223 NA-327 35.4 -88.4 Maastrichtian Maastrichtian j-j-h gb-a-3-$ 314 NA-328 39.7 -75.3 Campanian? Campanian? S-i-h gb-b-3-S 248 NA-329 38.9 -101.1 Campanian? Campanian? S-i-h gc-c-2-S 248 NA-33 68.6 -156.9 Camian Norian S-a-c a-C-3-F 290 NA-330 38.9 -101.1 Coniacian Santonian S-i-h gc-c-2-S 248 NA-331 32.4 -87.5 Campanian Campanian S-i-h gb-c-3-S 248 NA-332 40.3 -74.3 Maastrichtian Maastrichtian j-j-h gb-b-3-$ 248 NA-333 44.1 -99.6 Maastrichtian Maastrichtian j-j-h g-c-3-S 248 NA-334 40.4 -74.0 Maastrichtian Maastrichtian j-j-h gb-b-3-S 248 NA-335 40.3 -74.2 Maastrichtian Maastrichtian j-j-h gb-b-3-S 248 NA-336 25.7 -100.3 Turonian Campanian S-i-h gb-c-3-S 248; 251 NA-337 40.0 -74.7 Maastrichtian Maastrichtian j-j-h gb-b-3-S 248 NA-338 32.4 -87.3 Campanian Campanian S-i-h gb-b-2-$ 248;299 NA-339 33.6 -93.8 Campanian Campanian S-i-h gb-b-3-S 248; 251 NA-34 68.7 -158.4 Camian Norian S-a-c a-C-3-F 290 NA-340 43.4 -103.0 Campanian Campanian S-i-h gc-c-3-S 248 NA-341 30.3 -97.7 Campanian Campanian S-i-h gc-b-2-S 248;251 NA-342 37.2 -120.8 Campanian Maastrichtian S-i-h gd-d-2-S 248 NA-343 33.5 -88.4 Campanian Campanian S-i-h gb-b-2-S 248; 251 NA-344 38.8 -98.8 Campanian Campanian S-i-h gd-c-2-$ 248 NA-345 38.9 -101.2 Campanian Campanian S-i-h gb-c-2-S 248 NA-346 33.3 -88.1 Campanian Campanian S-i-h gd-d-3-S 248 NA-347 38.9 -101.1 Coniacian Campanian S-i-h gc-c-3-$ 248 NA-348 39.0 -99.9 Coniacian Campanian S-i-h gb-c-3-$ 248 NA-349 44.4 -100.3 Campanian Campanian S-i-h gc-c-2-S 248 NA-35 69.6 -144.9 Camian Norian S-a-c a-C-3-F 290 NA-350 39.9 -74.6 Maastrichtian Maastrichtian j-j-h gc-c-3-S 248 NA-351 38.9 -101.7 Campanian Campanian S-i-h gc-c-3-$ 248 NA-352 43.7 -103.4 Maastrichtian Maastrichtian j-j-h gd-b-3-S 248 NA-353 43.5 -102.7 Maastrichtian Maastrichtian j-j-h gb-b-3-S 248 312

TABLE G.l Marine Reptile Database Localities Age Range Lat Lon Maximum Minimum Interim Reliability Reference

NA-354 39.8 -75.2 Maastrichtian Maastrichtian j-j-h gb-b-3-S 248 NA-355 36.7 -119.8 Campanian Maastrichtian S-i-h gd-c-3-S 248 NA-356 40.1 -74.5 Maastrichtian Maastrichtian g-b-3-S 248 NA-357 38.8 -100.8 Coniacian Santonian S-i-h gc-c-3-$ M.J. Everhart. personal communication 1997; 248 NA-358 39,3 -97.8 Coniacian Campanian $-i-h gb-c-3-S 248 NA-359 42.8 -97.9 Coniacian Campanian S-i-h g-c-3-$ 248:319 NA-360 31.5 -97.1 Cenomanian Turonian S-i-h gc-c-3-S 248; 251 NA-361 49.0 -112.0 Upper Cretaceous Upper Cretaceous S-$-h gc-c-3-S 248 NA-362 46.8 -100.8 Upper Cretaceous Upper Cretaceous S-$-h gc-c-3-S 248 NA-363 47.7 -109.6 Upper Cretaceous Upper Cretaceous S-S-h gd-c-3-$ 248 NA-364 33.8 -93.8 Campanian Campanian S-i-h gc-b-3-S 248; 251 NA-365 32.9 -88.1 Campanian Campanian S-i-h gb-c-3-S 248 NA-366 32.8 -87.9 Campanian Campanian S-i-h gb-c-3-S 248 NA-367 33.5 -88.5 Campanian Campanian S-i-h g-b-2-S 248; 251 NA-368 32.8 -88.5 Campanian Campanian S-i-h gb-c-3-S 248 NA-369 29.2 -100.4 Coniacian Santonian S-i-h gb-c-3-$ 248; 251 NA-370 30.2 -97.6 Maastrichtian Maastrichtian j-j-h gb-b-3-S 248; 251 NA-371 33.1 -96.5 Campanian Campanian S-i-h gb-b-3-$ 248; 251 NA-372 33.4 -93.4 Santonian Santonian S-i-h gb-b-3-S 248; 251 NA-373 43.6 -109.6 Upper Cretaceous Upper Cretaceous S-S-h gd-c-3-$ 248 NA-374 43.0 -98.6 Campanian Maastrichtian S-i-h gb-c-3-$ 248 NA-375 43.3 -103.8 Campanian Campanian S-i-h gc-b-2-S 248 NA-376 38.9 -101.8 Campanian Campanian S-i-h gc-b-2-S 248 NA-377 43.0 -104.5 Campanian Campanian S-i-h gc-b-2-S 248 NA-378 43.0 -103.9 Campanian Campanian S-i-h gd-b-2-S 248 NA-379 42.9 -97.4 Campanian Campanian S-i-h gb-b-2-$ 248 NA-380 38.9 -101.8 Campanian Campanian S-i-h gc-b-2-S 248 NA-381 43.2 -104.3 Campanian Campanian S-i-h gb-b-2-S 248 NA-382 43.7 -103,5 Campanian Campanian S-i-h gb-b-2-S 248 NA-383 38.9 -101.6 Campanian Campanian S-i-h gc-b-2-$ 248 NA-384 43.4 -104.2 Campanian Campanian S-i-h gc-b-2-S 248 NA-385 43.2 -103.8 Campanian Campanian S-i-h gb-b-2-S 248 NA-386 43.7 -103.2 Campanian Campanian S-i-h gb-b-2-S 248 NA-387 37.6 -104.8 Campanian Campanian S-i-h gc-b-2-S 248 NA-388 45.5 -107.5 Santonian Campanian S-i-h gd-c-2-$ 248 NA-389 38.9 -101.6 Campanian Campanian S-i-h gb-b-2-S 248 NA-390 44.4 -100.0 Maastrichtian Maastrichtian j-j-h gb-b-2-S 248 NA-39I 45.9 -98.7 Maastrichtian Maastrichtian j-j-h gb-b-2-$ 248 NA-392 47.3 -109.1 Maastrichtian Maastrichtian j-j-h gc-b-2-S 248 NA-393 43.7 -103.5 Maastrichtian Maastrichtian j-j-h gb-b-2-S 248 NA-394 43.0 -97.1 Coniacian Campanian S-i-h gb-c-2-S 269 NA-42 40.3 -118.1 Anisian Anisian S-a-b c-S-?-$ 192; 274 NA-43 40.5 -109.5 Callovian Oxfordian S-d-e C-D-2-S 68 NA-467 37.2 -121.0 Campanian Maastrichtian S-i-h gc-c-2-$ 50 NA-468 36.7 -120.7 Campanian Maastrichtian M-h gb-c-2-S 50 NA-469 37.5 -121.1 Campanian Maastrichtian S-i-h gb-c-2-S 50 NA-470 36.8 -120.8 Campanian Maastrichtian S-i-h gb-c-3-S 50 NA-47I 36.1 -120.4 Campanian Maastrichtian S-i-h gc-c-3-S 50 NA-472 36.7 -120.7 Campanian Maastrichtian S-i-h gb-c-2-S 50 NA-493 32.8 -87.9 Campanian Campanian S-i-h gb-b-l-S 33;25I NA-494 32.8 -96.7 Santonian Santonian S-i-h a-b-3-S 298 NA-495 31.5 -96.9 Campanian Campanian S-i-h gb-c-3-$ 298; 251 NA-496 33.4 -96.9 Campanian Campanian S-i-h gb-b-3-S 298; 251 NA-497 33.2 -96.4 Campanian Campanian S-i-h gb-b-3-$ 298; 251 NA-498 33.5 -95.8 Campanian Campanian S-i-h gb-b-3-S 298; 251 NA-499 32.8 -96.8 Coniacian Santonian ^i-h a-c-3-S 298; 251 NA-5 74.0 -120.0 Maastrichtian Maastrichtian j-j-h b-B-2-DE 233 NA-500 32.3 -96.7 Campanian Campanian S-i-h a-b-2-S 298 313

TABLE G.l Marine Reptile Database Localities Age Range Lat Lon Maximum Minimum Interim Reliability Reference

NA-50I 32.2 -96.5 Campanian Campanian $-i-h a-b-2-S 298 NA-502 32.7 -96.3 Maastrichtian Maastrichtian j-j-h gb-b-3-S 298;251 NA-503 33.8 -96.7 Albian Albian $-h-g gb-b-3-S 277 NA-504 33.3 -97.2 Albian Albian $-h-g gb-b-3-S 277 NA-505 33,8 -96.7 Albian Albian $-h-g gb-b-3-S 277 NA-506 30.3 -97.7 Albian Cenomanian $-h-g gc-c-3-$ 277 NA-510 44.6 -120.2 Albian Albian $-h-g gb-b-7-S 194:217 NA-5II 44.8 -117.2 Norian Norian b-b-c gb-b-l-df 217 NA-5I7 22.6 -83.7 Oxfordian Oxfordian $-e-f b-b-3-S 309;130 NA-S2 44.4 -105.1 Albian Albian $-h-g d-b-2-F 202;17 NA-523 51.1 -100.0 Campanian Maastrichtian S-i-h gb-c-2-cf 16; 207 NA-525 49.3 -98.2 Upper Cretaceous Upper Crctaceous $-$-h gb-c-2-ef 16 NA-526 53.2 -102.6 Campanian Maastrichtian $-l-h ge-c-3-cf 16; 190 NA-529 39.4 -98.8 Turonian? Turonian? $-i-h gc-c-3-$ 309 NA-531 39.8 -75.2 Turonian Turonian S-i-h gb-c-3-$ 309 NA-578 39.0 -99.0 Cenomanian Turonian S-h-h gb-c-3-S 251;56 NA-579 42.8 -107.4 Cenomanian Cenomanian h-h-h gb-b-2-S 56 NA-580 30.3 -97.7 Turonian Turonian S-i-h gb-d-3-S 56 NA-581 39.4 -98.8 Cenomanian Turonian S-h-h gc-c-2-S 251; 56 NA-583 50.8 -111.5 Campanian? Campanian? S-i-h gc-c-3-S 253 NA-589 36.5 -104.9 Campanian Campanian S-i-h gb-b-4-df 268 NA-590 40.9 -113.0 Maastrichtian Maastrichtian j-j-h a-b-3-S 279; 144 NA-591 57.0 -122.9 Norian Norian b-b-c gc-b-3-f 283 NA-594 44.0 •104.4 Albian Albian S-h-g gb-b-2-$ 242; 173 NA-595 32.6 -87.7 Campanian Campanian S-i-h gb-b-2-$ 249;251 NA-596 32.8 -88.0 Campanian Campanian S-i-h gb-b-2-S 249;251 NA-597 32.9 -88.1 Campanian Campanian S-l-h gb-b-2-S 249;251 NA-598 39.5 -100.5 Coniacian Campanian S-i-h gd-b-3-S 307;229 NA-599 40.4 -74.2 Upper Cretaccous Upper Cretaceous S-S-h gc-c-3-S 307; 229 NA-6 74.0 -120.0 Maastrichtian Maastrichtian j-j-h b-B-2-DE 233 NA-600 30.8 -112.6 Camian Camian S-a-c a-b-l-c 49; 155 NA-601 54.5 -120.8 Smithian Spathian S-a-a b-c-2-f 47 NA-602 54.5 -120.7 Anisian Ladinian S-a-b a-e-2-f 47 NA-62 40.1 -117.6 Anisian Anisian S-a-b ge-A-2-F 259 NA-629 34.3 -117.5 Upper Cretaceous Upper Cretaceous S-S-h gb-c-3-S 158 NA-63 40.4 -117.5 Anisian Anisian S-a-b gd-A-2-F 259 NA-630 36.4 -110.4 Cenomanian Turonian S-h-h c-b-2-ef 154 NA-664 35.4 -105.9 Coniacian? Coniacian? i-i-h gc-c-2-df 156 NA-670 32.6 -87.3 San Ionian Santonian S-i-h gb-b-2-S 270 NA-671 32.5 -86.2 Santonian Campanian S-i-h gb-c-3-$ 270 NA-672 32.7 -88.1 Maastrichtian? Maastrichtian? j-j-h gb-c-2-S 270 NA-675 43.0 -106.8 Oxfordian? Oxfordian? S-d-e gd-b-2-S 89 NA-676 17.7 -97.9 Neocomian Neocomian S-f-g b-c-5-S 96 NA-682 37.7 -121.4 Upper Jurassic Lower Cretaceous S-S-f gb-d-6-$ 51 NA-683 37.5 -121.1 Upper Jurassic Lower Cretaceous S-S-f gb-d-6-S 51 NA-684 46.3 -106.7 Campanian Campanian s-i-h gb-b-3-$ 77 NA-685 37.3 -107.1 Campanian? Campanian? S-i-h gb-c-3-S 140 NA-686 53.9 -1332 Sinemurian Sinemurian S-c-d d-c-3-f 86 NA-692 54.5 -120.7 Anisian Ladinian S-a-b b-c-3-f 210 NA-693 31.6 -912 Cenomanian Cenomanian h-h-h gb-b-3-S 297 NA-694 35.1 -120.3 Tithonian Tithonian e-e-f gb-b-7-S 308 NA-695 32.8 -96.8 Turonian Turonian S-i-h gc-b-2-S 311 NA-696 40.1 -74.5 Maastrichtian Maastrichtian j-j-h gb-a-3-S 10 NA-697 50.6 -107.8 Campanian Campanian S-i-h gb-a-3-S 10 NA-698 40.3 -74.2 Maastrichtian Maastrichtian j-j-h gb-b-3-S 45; 102 NA-699 38.8 -76.9 Maastrichtian Maasuichtian j-j-h gc-a-3-S 12 NA-7 75.7 -119.5 Maastrichtian Maastrichtian j-j-h b-B-2-DE 233 NA-700 32.2 -84.9 Campanian Campanian S-i-h gb-b-3-d 266 NA-70I 32.1 -86.8 Maastrichtian Maastrichtian j-j-h gb-b-l-df 43 314

TABLE G.l Marine Reptile Database Localities ARC Range Lat Lon Maximum Minimum |nt^^ l^eliability Reference

NA-71 42.6 -124.4 Tithonian Tithonian e-e-f ge-A-2-F 53 NA-710 50.6 -107.4 Upper Campanian Lower $-i-h gb-c-3-f 300 Maastrichtian NA-711 39.4 -109.0 Upper Cretaceous Upper Cretaceous $-$-h gb-c-3-S 58 NA-712 34.6 -78.6 Campanian Campanian S-i-h ge-a-3-$ 195 NA-714 77.2 -95.9 Anisian Camian S-a-b gc-b-3-S 252 NA-716 75.2 -110.0 Aptian Albian $-g-g c-b-3-S 301;251;252 NA-719 57.9 -156.0 Kimmeridgian Portlandian $-c-r gb-c-3-$ 161 NA-73 40.8 -122.1 Camian Camian $-a-c gd-C-2-$ 192;48 NA-734 77.2 -95.9 Upper Triassic Upper Triassic $-$-c gc-c-6-S 48 NA-736 61.6 -143.2 Norian Norian b-b-c b-b-3-S 48 NA-737 50.6 -127.5 Norian Norian b-b-c gb-b-3-S 48 NA-74 39.6 -117,5 Anisian Anisian S-a-b gd-b-4-$ 192;48 NA-75 32.8 -97.4 Albian Albian $-h-g g*-B-6-S 191 NA-757 42.5 -111.4 Smithian Spathian S-a-a gb-b-3-S 164 NA-8 75.7 -119.5 Maastrichtian Maastrichtian j-j-h b-B-2-DE 233 NA-817 77.6 -111.0 Aptian Albian $-g-g C-C-3-S 30I;25I NA-818 39.8 -98.2 Coniacian Coniacian i-i-h gb-b-2-$ 309;251 NA-819 48.6 -98.0 Coniacian Santonian $-i-h gb-c-3-S 248;251 NA-82 49.6 -114.5 Sinemurian Sinemurian $-c-d gd»-C-3-S 205 NA-820 38.9 -101.1 Campanian Campanian S-i-h gc-a-2-dr [M.J. Everhart. personal communication 1997] NA-821 39.4 -99.9 Coniacian Santonian S-i-h gc-c-2-S [M.J. Everhart. personal communication 1997| NA-822 39.8 -99.4 Coniacian Santonian S-i-h gc-c-2-S [M.J. Everhart, personal communication 1997) NA-823 39.4 -99.3 Coniacian Santonian S-i-h gc-c-2-S [M.J. Everhart. personal communication 1997] NA-824 38.9 -99.3 Coniacian Santonian S-i-h gc-c-2-S [M.J. Everhart. personal communication 1997) NA-850 32.8 -88.0 Maastrichtian Maastrichtian j-j-h gb-b-2-S 299 NA-851 32.7 -87.6 Campanian Campanian S-i-h gb-c-2-S 299 NA.852 41.4 •106.6 Upper Jurassic Upper Jurassic S-S-c gc-c-3-S 112 NA-853 43.6 -106.8 Upper Jurassic Upper Jurassic S-S-e gc-c-3-S 112 NA-854 39.3 -103.1 Campanian Maastrichtian S-i-h gb-c-3-S 140 NA-855 37.2 -104.5 Campanian Maastrichtian S-i-h gb-c-3-S 140 NA-8S9 39.5 -75.7 Campanian Campanian S-i-h ab-a-3-S 14 NA-860 39.1 -75.6 Campanian Campanian S-i-h b-a-3-S 14 NA-864 42.6 -96.5 Cenomanian Cenomanian h-h-h b-b-3-S 319;251 NA-865 35.1 -107.1 Santonian Santonian S-i-h b-b-2-S 224;251 NA-866 36.8 -107.2 Campanian Campanian S-i-h gb-b-6-S 157:251 NA-867 32.8 -108.5 Upper Cretaceous Upper Cretaceous S-S-h gb-c-3-S 157:251 NA-868 33.2 -107.1 Upper Cretaceous Upper Cretaceous S-S-h gb-c-6-S 157 NA-869 36.5 -104.9 Upper Cretaceous Upper Cretaceous S-S-h gb-c-3-S 157;251 NA-870 36.6 -104.8 Campanian Maastrichtian S-i-h gb-c-3-$ 157;251 NA-872 45.1 -96.3 Turonian Turonian S-i-h gc-b-6-S 278; 251 NA-873 45.2 -96.6 Turanian Turonian S-i-h gb-b-6-S 278;251 NA-874 76.2 -119.2 Tithonian Tithonian e-«-f c-b-2-f 301 NA-875 76.2 -119.2 Tithonian Tithonian e-e-f c-b-3-f 301 NA-876 77.7 -112.6 Oxfordian Kimmeridgian S-e-f c-c-3-f 301 NA-878 33.8 -93.9 Campanian Campanian S-i-h gb-b-3-S 307;251 NA-879 33.8 -94.1 Santonian Santonian S-i-h gb-b-3-S 307; 251 NA-881 29.3 -103.6 Campanian Campanian S-i-h gb-b-3-S 320;251 NA-882 43.6 -104.1 Albian Albian S-h-g gc-b-2-S 202;251 NA-883 40.5 -74.4 Cenomanian Cenomanian h-h-h gb-b-3-S 101 NA-884 40.0 -75.0 Campanian Campanian S-i-h gb-b-3-S 101 NA-885 40.1 -74.6 Campanian Campanian S-i-h gb-b-3-$ 102 NA-886 40.1 -74.6 Maastrichtian Maastrichtian j-j-h gb-b-3-S 102 NA-894 38.5 -118.1 Norian Norian b-b-c b-b-2-def 145 315

TABLE G.l Marine Reptile Database Localities Age Range

Lat Lon Maximum Minimum Interim Reliability Reference

NA-898 42.7 -121.2 Campanian Campanian S-i-h ac-a-2-df 124 NA-9 69.0 -129.0 Campanian Campanian $-i-h C-B-3-DE 233 SA-28 -39.2 -69.7 Tithonian Tithonian e-c-f a-A-2-F 94; 106 SA-280 9.9 -66.4 Cenomanian Turonian $-h-h g-C-$-$ 307;229 SA-281 -36.6 -73.1 Maastrichtian Maastrichtian j-j-h a-b-$-$ 229; 107;60 SA-282 -43.4 -69.2 Maastrichtian Maastrichtian j-j-h gb-B-$-$ 229 SA-283 -36.7 -73.1 Maastrichtian Maastrichtian j-j-h gb-b-$-$ 229; 107;60 SA-284 -28.1 -65.9 Lower Jurassic Lower Jurassic $-S-d gd-C-$-$ 229 SA-29 -38.7 -70.0 Tithonian Tithonian e-e-f d-A-2-F 106 SA-30 -38.0 -70.0 Tithonian Tithonian e-e-f d-A-2-F 106 SA-31 -36.0 -70.0 Tithonian Tithonian e-e-f d-B-2-F 106 SA-326 8.6 -70.2 Upper Cretaceous Upper Cretaceous S-$-h ge-c-3-f 231 SA-40 -39.3 -70.3 Bajocian Bajocian $-d-e d-A-2-F 93 SA-44 -26.1 -69.3 Upper Triassic Upper Triassic $-$-c a-C-l-DEF 287 SA-492 -35.5 -69.6 Tithonian Tithonian e-e-f ge-c-6-J 246; 37 SA-50 -71.4 -47.9 Neocomian Neocomian $-f-g d-C-6-F 17 SA-508 4.6 -74.9 Coniacian Coniacian i-i-h gc-b-3-f 123 SA-518 -28.0 -70.1 Lower Jurassic Lower Jurassic $-$-d a-c-6-$ 309; 66; 60 SA-519 5.6 -73.6 Aptian Aptian g-g-g ge-b-3-f 309;119 SA-534 -34.0 -69.7 Callovian Callovian d-d-e gc-b-4-$ 37; 108 SA-536 -39.4 -70.1 Bajocian Bajocian $-d-c gc-b-4-$ 37; 108 SA-537 -8.8 -37.7 Maastrichtian Maastrichtian j-j-h gd-c-3-$ 37; 59 SA-585 -8.8 -74.5 Cenomanian Cenomanian h-h-h gb-c-3-$ 19 SA-680 -39.3 -70.3 Callovian Callovian d-d-e gb-a-2-def 109 SA-687 -5.5 -78.6 Cenomanian Cenomanian h-h-h gb-b-3-df 131 SA-718 10.4 -72.2 Aptian Aptian g-g-g gc-b-2-f 198 SA-72I -32.7 -70.7 Toarcian Toarcian $-c-d c-a-4-f 108 SA-722 -36.9 -69.7 Tithonian Tithonian e-c-f gb-b-4-f 108 SA-723 -37.2 -70.1 Tithonian Tithonian e-e-f d-b-4-f 108 SA-724 -37.2 -70.4 Tithonian Tithonian e-c-f gb-b-4-r 108 SA-725 -34.1 -69.7 Berriasian Berriasian $-f-g c-b-4-f 108 SA-726 -48.9 -72.7 Barremian Aptian S-S-g gc-b-4-f 108 SA-758 -10.0 -74.9 Santonian Santonian $-i-h a-b-3-ef 46 SA-77 -32.0 -68.6 Lower Jurassic Lower Jurassic $-$-d gd-C-4-$ 245 SA-78 -32.0 -68.6 Bajocian Bajocian $-

TABLE G.l Marine Reptile Database Localities Age Range Loc Mso IP Lat Lon Maximum Minimum Reliability Rererence

SA-893 -35.6 -72.6 Maastrichtian Maastrichtian j-j-h a-b-3-$ 60 TABLE G.2 Marine Reptile Database Taxa

Loc Map MMR Order Family (icnus Species Cuniplctc fcscr- Rgfefjnc,. ID Interval ID • • vntfr%n

AF-239 h-h-h MR-406 Plesiosauria indei. indet. indel. c 0 229 AF-239 h-h-h MR-405 Plesiosauria Polycolylidac indet. indel. e 0 229 AH-239 h-h-h MR-404 Plesiosauria Polycolylidae indel. indel. b/c 0 229 •AF-241 $-r-g MR-408 Plesiosauria Pliosauridac I.eplocleidus capcnsis b/c/i 0 309; 229; 75 AF-3M $-i-h MR-1404 Plesiosauria t:lasmosauridae indel. indel. d/z 0 313 AF-314 $-i-h MR-530 Squamala Mosasauridae I.eiodon indel. d 1 90 AF-314 $-i-h MR-528 Squamaia Mosasauridac Mosasaurus indel. d 1 90 AF-314 S-l-h MR-529 Squamala Mosasauridae Platccarpus indel. d 1 90 AF-315 h-h-h MR-532 Plesiosauria indet. indel. indel. i. 0 90 AF-316 j-j-h MR-533 Squaniala Mosasauridac Goronyosaurus nigeriensis c 3 281 AF-317 j-j-h MR-534 Squamala Mosasauridae Goronyosaurus nigeriensis b/c/d/c/h 4 9; 281 AF-317 j-j-h MR-536 Squamala Mosasauridae Halisaurus indel. e 3 9 AF-317 j-j-h MR-535 Squamala Mosasauridae indel. indel. e 3 9 AF-37 $-l-h MR-63 Plesiosauria indel. indel. indel. i 4 149 AF-37 $-i-h MR-62 Squamala Mosasauridac Plaiccarpus bocagei b/c/d/e 4/5 149 AF-37 S-l-h MR-632 Squamala Mosasauridae Tylosaurus iembeensis b/c/d/e 6 293; 147 AF-38 j-j-h MR-71 Squamala Mosasauridae indel. indel. d 2 148 AF-38 j-j-h MR-67 Squamala Mosasauridae Mosasaurus cf. M. holTmanni d 2 148 AF-38 j-j-h MR-66 Squamala Mosasauridae Mosasaurus cf. M. lemonnieri d 2 148 AF-38 j-j-h MR-69 Squamala Mosasauridac Mosasaurus sp. e 2 148 AF-38 j-j-h MR-64 Squamala Mosasauridac I'lioplalecarpus sp. d 2 148 AF-38 j-j-h MR-65 Squamala Mosasauridae Prognalhodon cf. P. giganteus d 2 148 AF-39 j-j-h MR-68 Squamala Mosasauridae Halisaurus sp. c 2 148 AF-39 j-j-h MR-70 Squamala Mosasauridae Mosasaurus sp. e 2 148 AF-398 j-j-h MR-625 Plesiosauria indel. indel. indel. d/e 0 293 AF-398 j-j-h MR-637 Squamala Mosasauridae indet. indel. d 0 293 AF-399 j-j-h MR-626 Plesiosauria indel. indet. indel. d/e 0 293 AF-399 j-j-h MR-638 Squamala Mosasauridae indet. indel. d/e 0 293 AF-400 j-j-h MR-627 Plesiosauria indet. indet. indel. c 0 293 AF-400 j-j-h MR-639 Squamala Mosasauridae indet. indel. d/e 0 293 AF-401 j-j-h MR-628 Plesiosauria indel. indet. indci. e 0 293 AF-401 j-j-h MR-6SI Squamala Mosasauridac Globidens indet. d 0 293 AF-401 j-j-h MR-643 Squamala Mosasauridae indet. indel. d/e/z 0 293 AF-402 j-j-h MR-629 Plesiosauria indet. indet. indel. d 0 293 AF-402 j-j-h MR-647 Squamala Mosasauridae indci. indel. d 0 293 AF-403 j-j-h MR-630 Plesiosauria indel. indet. indel. d 0 293 AF-403 j-j-h MR-649 Squamala Mosasauridae indet. indel. d 0 293 AF-404 j-j-h MR-648 Squamala Mosasauridae indel. indel. d 0 293 TABLE G.2 Marine Reptile Database Taxa

l-OC Map MMR Order Family Genus Spccies Compleic .. Rcfercncc ID nlerval ID ' ' valion

Ar-405 h MR-633 Squamala Mosasauridae indet. indet. d/c 0 293 AF-406 h MR-634 Squamala Mosasauridae indet. indel. z 0 293 A1--407 h MR-635 Squamata Mosasauridae indet. indet. d 0 293 AF-408 h MR-650 Squamala Mosasauridae Cilobidens indet. d 0 293 AF-408 h MR-636 Squamala Mosasauridae indet. indet. d 0 293 AI--409 h MR-640 Squamata Mosasauridae indet. indel. d/e 0 293 AF-410 -h MR-641 Squamata Mosasauridae indet. indel. d/z 0 293 AF-4II -h MR-642 Squamata Mosasauridae indet. indel. i/z 0 293 AF-412 h MR-644 Squamala Mosasauridae indet. indel. d//7h 0 293 AF-413 h MR-645 Squamata Mosasauridae indet. indet. d 0 293 AF-414 h MR-646 Squamata Mosasauridae indet. indet. d/z 0 293 AF-447 h MR-1286 Squamata Mosasauridae Goronyosaurus sp. b/c/d 3 146 AF-447 h MR-1289 Squamata Mosasauridae llalisaurus sp e 3 146 AF-447 h MR-1287 Squamata Mosasauridae Igdamaiiosaurus aegyptlacus c/d 3 146 AF-447 -h MR-1292 Squamata Mosasauridae indet. indet. b/c/d/crti/i 3 146 AF-447 -h MR-I29I Squamata Mosasauridae Mosasaurus M. holTmanni d 2 146 AF-447 h MR-1288 Squamata Mosasauridae Plalecarpus bocagei c 3 146;149 AF-447 h MR-704 Squamala Mosasauridae Platecarpus bocagci c 5 149 AF-447 h MR-1290 Squamala Mosasauridae Plioplalecarpus sp. c 3 146 AF-544 h MR-962 IMesiosauria lilasmosauridae indet. indel. e 6 AF-544 h MR-964 Squamala Mosasauridae t.eiodon cf L. anceps d 4 6 AF-545 h MR-963 Plesiosauria niasmosauridae indet. indel. e 5 6 AF-545 -h MR-982 Squamata Mosasauridae Igdanianosaurus aegypliacus d 4 6; 146 AF-545 -h MR-976 Squamata Mosasauridae indet. indel. d/c 4 6 AF-545 h MR-970 Squamata Mosasauridae indet. indel. d 4 6; 146 AF-545 h MR-965 Squamala Mosasauridae Leiodon cf. I., anceps d 4 6 AF-546 h MR-983 Squamata Mosasauridae Igdamanosauius aegypliacus d 4 6; 146 AF-546 -h MR-977 Squamala Mosasauridae indet. indet. d 4 6 AF-546 h MR-971 Squamata Mosasauridae indet. indet. d 4 6; 146 AF-546 h MR-966 Squamata Mosasauridae I.eiodon cl'. 1.. anceps d 4 6 AF-548 -h MR-985 Plesiosauria Ulosmosauridae indet. indet. d 4 6 AF-548 -h MR-973 Squamata Mosasauridae indet. indet. d 4 6; 146 AF-548 h MR-978 Squamata Mosasauridae indet. indet. d 4 6 AF-548 h MR-968 Squamala Mosasauridae I.eiodon cf. 1.. anceps d 4 6 AF-549 h MR-986 Plesiosauria Flasmosauridae indet. indel. d 4 6 AF-549 h MR-979 Squamala Mosasauridae indet. indel. d 4 6 AF-550 h MR-987 Plesiosauria lilasmosauridae indet. indel. d 4 6 AF-550 h MR-984 Squamala Mosasauridae Igdanianosaurus aegypliacus d 4 6; 146 TABLE G.2 Marine Reptile Database Taxa

Luc Map MMR Order Family Genus Specics Coniplcic . Rcfcrence ID Interval ID ' ' valion

AI--550 j-j-h MR-974 Squaniala Mosasauridae indet. indet. d 4 6; 146 AF-550 j-j-h MR-981 Squamala Mosasauridae I.eiodon cf. 1.. anceps d 4 6 AF-551 j-j-h MR-988 Plesiosauria HIasmosauridae indet. indet. d 4 6 Ar-551 j-j-h MR-980 Squamata Mosasauridae indet. indet. d 4 6 AI--55I j-j-h MR-975 Squamata Mosasauridae indet. indet. d 4 6; 146 AF-551 j-j-h MR-969 Squamala Mosasauridae I.eiodon cf 1.. anceps d 4 6 AF-552 j-j-h MR-989 Plesiosauria HIasmosauridae indet. indet. d 4 6 AF-673 $-a-b MR-1229 Plesiosauria Pistosauridae indet. indet. e 4 120;240 AF-691 $-i-h MR-1262 Plesiosauria HIasmosauridae indet. indet. e 4 28 AF-732 $-$-g MR-1346 Plesiosauria Plesiosauridae cl'. Plesiosaunis sp. d/z 2 265 AI--760 c-c-f MR-1397 Ichthyosauria Ichthyusauridae Brachypterygius sp. b 5 95 AF-760 e-«-f MR-1398 Ichthyosauria indet. indet. indet. 1 6 95 AF-761 j-j-h MR-1399 Plesiosauria HIasmosauridae indet. indet. e 4 313 AF-761 j-j-h MR-1400 Squamala Mosasauridae indet. indet. t 0 313 AF-83 $-f-g MR-162 Plesiosauria Plesiosauridae Plesiosaunis sp. e 3 126 AF-861 $-i-h MR-1544 Plesiosauria indet. indet. indet. d 0 235 AI--862 S-i-h MR-1545 Squamata Mosasauridae indet. indet. d 0 235 AF-896 j-j-h MR-1608 Squamata Mosasauridae Pluridens walkeri c/d 4 152 AN-628 $-i-h MR-1301 Plesiosauria Cryptoclididae Mortumeria seyniourensis b/c/d/e 6 62; 61 AN-628 $-i-h MR-1302 Plesiosauria HIasmosauridae indet. indet. e/h/i 5 62 AN-628 S-i-h MR-1131 Plesiosauria indet. indet. indet. b/c/e/h/i 6 64; 63 AN-628 $-i-h MR-1132 Squamata Mosasauridae indet. indet. b/e/i 6 64; 63 AN-841 S-i-h MR-1516 Plesiosauria indet. indet. indet. e/r 4 107 AN-842 S-i-h MR-1517 Plesiosauria indet. indet. indet. di 0 107 AS-2 S-$-b MR-2 Ichthyosauria indet. indet. indet. S 0 48 AS-212 S-S-h MR-368 Plesiosauria '.'Pliosauroidea indet. indet. e 0 309; 229 AS-212 $-S-h MR-361 Plesiosauria HIasmosauridae indet. indet. e/i 0 309; 229 AS-212 $-$-h MR-363 Plesiosauria HIasmosauridae indet. indet. e 0 309;229 AS-212 S-S-h MR-383 Plesiosauria indet. indet. indet. e 0 229 AS-212 S-S-h MR-371 Plesiosauria Pliosauroidea indet. indet. e 0 309; 229 AS-212 S-S-h MR-366 Plesiosauria Polycotylidae Pulycutylus cf. P. latipinnis c 0 229;57 AS-212 S-S-h MR-1194 Squamata Mosasauridae indet. indet. b 0 244 AS-213 S-S-h MR-367 Plesiosauria '.'Pliosauroidea indet. indet. e/i 0 309;229 AS-213 S-S-h MR-369 Plesiosauria '.'Pliosauroidea indet. indet. e 0 309; 229 AS-213 S-S-h MR-362 Plesiosauria I'Masmosauridae IHasmosaurus indet. e 0 309; 229 AS-213 S-S-h MR-370 Plesiosauria indet. indet. indet. e 0 229 AS-213 S-S-h MR-365 Plesiosauria Pliosauridae indet. indet. e 0 309; 229 AS-214 h-h-h MR-364 Plesiosauria HIasmosauridae indet. indet. e/i 0 309; 229 TABLE G.2 Marine Reptile Database Taxa

Loc Map MMR Preser­ Order Pamily Genus Species Complete Rcfercncc ID 1Interval ID vation

AS-215 h-h-h MR-372 Plesiusauria indel. indel. indel. c 0 309; 229 AS-215 h-h-h MR-373 Pleslosauria Polycoiylidae '.'Dolichorhynchop sp. e/i 0 309;229 S AS-216 h-h-h MR-374 Plesiosauria Pliosauridae I'olypiychodon inierruplus d 0 229 AS-217 $-S-h MR-375 Pleslosauria indel. indel. indel. c/i 0 229 AS-218 $-h-g MR-376 Pleslosauria indel. indet. indet. 1 0 229 AS-219 $-$-g MR-1436 Ichlhyosauria indel. indel. indet. e 0 204 AS-219 $-$-g MR-377 Pleslosauria indel. indel. indet. i 0 229 AS-220 e-e-f MR-378 Pleslosauria indel. indel. indet. e 0 229;40 AS-221 c-c-r MR-379 Pleslosauria indel. indel. indel. c/i 0 229 AS-222 c-c-f MR-380 Pleslosauria ?Cryploclididac Colymbosaurus sp. i 0 229 AS-223 e-c-f MR-381 Pleslosauria Cryploclididac Colymbosaurus? cf. C. irochanterius e 0 229 AS-223 e-«-f MR-382 Pleslosauria indel. indel. indel. e 0 229 AS-225 $-d-c MR-387 Pleslosauria lilasmosauridae Muraenosaurus sp. e 0 229 AS-226 d-d-c MR-388 Pleslosauria Elasmosauridae Muracnosaurus leedsii e 0 229 AS.227 d-d-c MR-389 Pleslosauria indel. indel. indet. e/i 0 229 AS-228 d-d-e MR-390 Pleslosauria Cryploclididae Crypioclidus? sp. e 0 229 AS-229 e-c-f MR-391 Pleslosauria Pliosauridae Liopleurodon cl'. L. macromerus d/e 0 229 AS-230 $-e-f MR-392 Plesiosauria Pliosauridae Pliosaurus brachyspondylus d/c 0 229 AS-231 $-c-f MR-393 Pleslosauria indel. indel. indet. b 0 229 AS-232 s-c-r MR-394 Plesiosauria indel. indel. indel. b 0 229 AS-233 $-e-f MR-395 Plesiosauria Pliosauridae Peloneusies philarchus e/i 0 229 AS-235 S-S-d MR-398 Plesiosauria indet. indet. indet. e 0 229 AS-236 $-$-h MR-399 Plesiosauria tlasmosauridae indel. indel. i 0 229 AS-237 $-e-r MR-401 Plesiosauria Pliosauridae indel. indel. d 0 229 AS-237 S-e-f MR-400 Plesiosauria Pliosauridae Pliosaurus cf. P. andrewsi d 0 229 AS-238 h-h-h MR-403 Plesiosauria indel. indel. indel. e 0 229 AS-247 $-e-f MR-414 Plesiosauria Pliosauridae Pclunuesles cf. P. philarchus c 0 229 AS.291 $-$-h MR-480 Plesiosauria lilasmosauridae indel. indel. e/h/i 0 309; 229 AS-291 $-$-h MR-II90 Squamala Mosasauridac indel. indel. d/c 5 244 AS-3 $-$-b MR-3 Ichlhyosauria Mixosaurldae Mixusaunis sp. S 0 48 AS-4 $-$-b MR-4 Ichlhyosauria Mixosauridae Mixosaurus sp. c 0 168; 48 AS-45 $-e-r MR-77 Ichlhyosauria Ichthyosauridae Ophlhalmosaurus sp. d/e/h 2/3 324 AS-49 $-h-g MR-109 Ichlhyosauria Plalyplerygidae Platyplcrygius kiprijanofTi b/d/e/i 6 204;17 AS-586 j-j-h MR-1060 Plesiosauria indel. indel. indet. d 2 262 AS.587 $-$-c MR-I06I Ichlhyosauria indel. indel. indel. d 2 215 AS-588 j-j-h MR-1403 Plesiosauria HIasmosauridae indel. indet. d 0 272;309;313 AS-588 j-j-h MR-1062 Squamala Mosasauridac Igdamanosaurus acgyptiacus d 4 8; 272;146 u> to o TABLE G.2 Marine Reptile Database Taxa

Loc Map MMR Order Family Ocnus Specics Complete . Reference ID 1Interval ID ' * vatinn

AS-S88 j-j-h MR-1602 Squamata Mosasauridae indet. indet. d 0 272 AS-S88 j-j-h MR-I60I Squaniata Mosasauridae Leiodon cf. M. anceps d 0 272; 20 AS-593 $-h-g MR-1071 Ichthyosauria Platypterygidae Platypterygius "campy lodon" e 0 173 AS-603 h-h-h MR-IIOI Plesiosauria HIasmosauridae indet. indet. e/f/g/h/i 6 214 AS-64 $-a-a MR-I3I Ichthyosauria incertac sedis lltatsusaurus hataii q 5/6 271 AS-646 $-8-a MR-II70 Ichthyosauria incertae sedis Chaohusaurus geishanensis s 0 323;20 AS-647 b-b-c MR-II7I Ichthyosauria incertae sedis Himalaysaurus tibetensis s 0 323; 20 AS-65 S-a-a MR-132 Ichthyosauria incertae sedis Lltatsusaurus hataii b/e/f/h 5/6 271 AS-653 b-b-e MR-II87 Ichthyosauria incertae sedis Tibetosaurus tingjiensis S 0 87; 48; 20 AS-655 $-$-h MR-II9I Squamata Mosasauridae indet. indet. h 5 244 AS-656 $-$-h MR-II92 Plesiosauria indet. indet. indet. e/i 0 244 AS-657 $-$-h MR-II93 Squamata Mosasauridae Prognathodon sp. a 6 244; 153 AS-658 $-$-h MR-l >95 Squamata Mosasauridae indet. indet. e 5 244 AS-659 j-j-h MR-1196 Squamata Mosasauridae indet. indet. z 0 244 AS-660 $-$g MR-l 197 Plesiosauria indet. indet. indet. z 0 244 AS-661 j-j-h MR-l 198 Squamata Mosasauridae indet. indet. b 5 244 AS-662 j-j-h MR-l 199 Squamata Mosasauridae indet. indet. d 4 244 AS-662 j-j-h MR-I45I Squamata Mosasauridae indet. indet. d 0 118 AS-67 S-a-b MR-134 Ichthyosauria Mixosauridae Mixosaurus maotaiensis e/l7g/h/i 6 325; 48 AS-677 $-$-a MR-1235 Ichthyosauria incertae sedis lliaisaurus chonglakmanii b/c/d/e/i 6 172 AS-678 S-l-h MR-1242 Squamata Mosasauridae Carinodens fraasi d 4 238; 298 AS-678 S-l-h MR-1240 Squamata Mosasauridae indet. indet. d 4 238 AS-678 S-i-h MR-l 239 Sqi'amata Mosasauridae Leiodon cf. L. anceps d 4 238 AS-678 S-i-h MR-1241 Squamata Mosasauridae Platecaipus ? sp. d 4 238 AS-688 $-e-f MR-1259 Plesiosauria Cryptoclididae indet. indet. e 6 24 AS-702 $-r-g MR-l 293 Ichthyosauria indet. Simbirskiasaurus birjukovi b/c/d/e 6 216 AS-706 $-i-h MR-1299 Plesiosauria i-lasmosauridac indet. indet. e/l7g/h/i 6 203 AS-706 $-i-h MR-1336 Squamata Mosasauridae indet. indet. S 0 213 AS-706 S-i-h MR-1334 Squamata Mosasauridae indet. indet. S 0 213 AS-706 S-i-h MR-l 335 Squamata Mosasauridae Plotosaurus? sp s 0 213 AS-708 S-S-e MR-l 304 Plesiosauria Pliosauridae? Yu/houpliosaurus chengjiangensis c/d/e/h/i 5/6 328 •AS-709 S-Sg MR-1305 Plesiosauria Pliosauridae Sinopliosaurus fusuiensis d 4 128 AS-717 s-i-h MR-I3I9 Plesiosauria indet. indet. indet. e/f/g/h/i 6 165 AS-727 j-j-h MR-1337 Squamata Mosasauridae indet. indet. S 0 213 AS-727 j-j-h MR-1338 Squamata Mosasauridae Mosasaurus sp. S 0 213 AS-728 j-j-h MR-1339 Squamata Mosasauridae indet. indet. s 0 213 AS-729 S-i-h MR-l 341 Squamata Mosasauridae indet. indet. s 0 213 AS-729 S-i-h MR-1340 Squamata Mosasauridae indet. indet. s 0 213 TABLE G.2 Marine Reptile Database Taxa

I.oc Mop MMR Order Family Genus Snecics Complete . Reference ID Interval ID * ' vnlmn

AS-730 S-i-h MR-1342 Squaiiiala Mosasauridae Mosasaurus? sp. $ 0 213 AS-738 b-b-c MR-1353 Ichthyosauria Mixosauridae Mixosaurus sp e 0 204 AS-738 b-b-c MR-1352 IchthyosBuria Shastasauridae indet. indet. e 0 204; 48 AS-739 $-a-c MR-1355 Ichthyosauria Shastosauridae ?Cymbospondylus sp. e/f 6 204; 48 AS-741 $-a-c MR-1357 Ichthyosauria Shastasauridae indet. indet. $ 0 48 AS-747 $-a-b MR-1367 Ichthyosauria Shastasauridae indet. indel. h 0 204;48 AS-748 $-a-b MR-1368 Ichthyosauria Shastasauridae Shastasaurus? sp. e/f 6 204; 48 AS-755 $-$-a MR-1391 Ichthyosauria indet. indet. indet. $ 0 48 AS-765 $-C-ll MR-1407 Ichthyosauria indet. indet. indel. S 0 204 AS-766 $-c-d MR-1408 Ichthyosauria indet. indet. indel. S 0 204 AS-767 $-d-e MR-1409 Ichthyosauria indet. indet. indel. s 0 204 AS-770 e-e-f MR-I4I2 Ichthyosauria indet. indet. indet. e 0 204 AS-771 $-$-r MR-1413 Ichthyosauria indet. indet. indet. e 0 204 AS-772 d-d-e MR-I4I4 Ichthyosauria indet. indel. indet. e 0 204 AS-773 e-e-f MR-I4IS Ichthyosauria indet. indet. indet. e 0 204 AS-774 e-e-f MR-1416 Ichthyosauria Ichthyosauri dae Ophthalmosaurus icenicus e/f/i 6 204 AS-777 e-e-f MR-1420 Ichthyosauria indet. indel. indet. e 0 204 AS-780 c-e-f MR-1578 Ichthyosauria Ichthyosauridae Ophthalmosaurus sp. a/q 6/7 137 AS-780 e-e-f MR-1423 Ichthyosauria indet. indet. indet. q 0 204 AS-780 e-e-f MR-1577 Pleslosauria Pliosauridae indel. indet. a/q 6/7 137 AS-781 c-e-f MR-1424 Ichthyosauria indet. indel. indet. e 6 204 AS-783 e-e-f MR-1426 Ichthyosauria indet. indel. indet. e 0 204 AS-784 e-e-f MR-1427 Ichthyosauria indet. indel. indet. b/c/d/e/h 0 204 AS-785 e-e-f MR-1428 Ichthyosauria indet. indet. indel. d/e 0 204 AS-786 f-f-8 MR-1429 Ichthyosauria indet. indel. indel. a 0 204 AS-787 f-f-g MR-1430 Ichthyosauria indet. indet. indel. b/e 0 204 AS-788 $-$-g MR-1431 Ichthyosauria indet. indet. indel. b 0 204 AS-789 $-$-g MR-1432 Ichthyosauria indet. indet. indet. e/i 6 2M AS-790 f-f-g MR-1433 Ichthyosauria indet. indel. indet. e 0 204 AS-791 $-$-g MR-1434 Ichthyosauria indet. indel. indel. b/c/d 0 204 AS-792 $-$•£ MR-1435 Ichthyosauria indel. indet. indel. e 0 204 AS-793 $-$-g MR-1437 Ichthyosauria indet. indel. indel. e 0 204 AS-794 $-h-g MR-1438 Ichthyosauria indet. indet. indel. b/c/d 0 204 AS-795 $-h-g MR-1439 Ichthyosauria Platyplerygidae Platypierygius kipriJanoITi b/d/e/f/i 5 204;19 AS-796 $-f-g MR-I4I9 Ichthyosauria Platypterygidae Plalypterygius kiprijanofTi b/c/e 0 204; 19 AS-797 $-h-g MR-1440 Ichthyosauria Platypterygidae Platypierygius kiprijanoITi d 0 204; 19 AS-798 h-h-h MR-I44I Ichthyosauria indet. indel. indet. d 0 204 AS-799 h-h-h MR-1442 Ichthyosauria indet. indet. indet. d/z 0 204 U) TABLE G.2 Marine Reptile Database Taxa

LJK Map MMK Order I-'amily Genus Species Complete . ' Reference ID 1Interval ID * * vaiion

AS-800 h-h-h MR-1443 Ichthyosauria Platypteo'gidae Platyptcrygius sp. c 0 204; 19 AS-801 h-h-h MR-1444 Ichthyosauria indei. indel. indel. e/r/i 0 204 AS-802 h-h-h MR-1445 Ichthyosauria indet. indel. indel. dm 0 204 AS-803 h-h-h MR-1446 Ichthyosauria indet. indel. indel. d/e/i 0 204 AS-804 h-h-h MR-1447 Ichthyosauria indet. indel. indel. eJ'i 0 204 AS-805 h-h-h MK-1448 Ichthyosauria indet. indel. indel. e 0 204 AS-806 $-$-g MR-1449 Ichthyosauria indet. indet. indel. die 0 204 AS-807 h-h-h MR-1450 Ichthyosauria indel. indel. indet. d/z 0 204 AS-808 $-$-h MR-1452 PIcsiosauria indel. indet. indel. S 0 132 AS-809 S-S-h MR-1453 Squamata Mosasauridae indet. indel. d/e/i 0 236 AS-810 S-i-h MR-1454 Squamata Mosasauridae indet. indel. e 0 139 AS-849 $-$-d MR-1526 Plesiosauria indet. indel. indel. $ 0 105 AS-887 $-$-d MR-1579 IMesiosauria indet. indet. indel. d 2 125 AS-888 $-h-g MR-1580 Plesiosauria indet. indel. indel. e 0 304 AS-895 $-$-g MR-1606 PIcsiosauria indet. indet. indel. /. 0 329 AS-91 $-i-h MR-171 Plesiosauria Polycotylidae Oeorgiasaunis penzensis b/c/d/em/i 6 219;220 •AS-92 $-$-<1 MR-172 Plesiosauria Pliosauridae Bislianopliosaurus youngi e/f/h/i 6 88 AS-93 $-$-f MR-173 Plesiosauria indet. indet. indel. e 2 44 AS-94 e-c-f MR-174 Plesiosauria Pliosauridae Simolestes indicus e 3 303;24 AS-94 e-c-f MR-402 Plesiosauria Pliosauridae Simolestes'.' indicus c 0 229; 303 AU-lOO B-8-6 MR-1483 Plesiosauria l-lasmosauridac indet. indet. $ 0 309;196 AU-lOO g-g-g MR-184 Plesiosauria indet. indet. indet. e 2 226;309 AU-lOO g-g-g MR-462 Plesiosauria Pliosauridae ?Kronosaurus sp. a 0 229; 296 AU-IOI g-g-g MR-185 PIcsiosauria lilasmosauridae indet. indel. e/f/h/i 4 226; 309 AU-102 $-h-g MR-189 Plesiosauria HIasmosauridae indet. indet. dOh 6 226 AU-102 $-h-g MR-186 Plesiosauria Elasmosauridae indel. indet. dh/z 6 226;309 AU-103 $-h-g MR-1486 Ichthyosauria indet. indet. indet. S 0 196 AU-103 $-h-g MR-187 Plesiosauria HIasmosauridae indel. indet. e 5 226; 309 AD-104 g-g-g MR-190 Plesiosauria HIasmosauridae indel. indel. d/. 3 226 AU-105 $-h-g MR-I9I Plesiosauria HIasmosauridae indel. indel. e/l7h/i/z 5 226 AU-285 $-$-h MR-464 Plesiosauria HIasmosauridae indel. indet. e^/i 0 229 AU-285 S-$-h MR-465 Plesiosauria HIasmosauridae indet. indet. e 0 229 AU-285 $-$-h MR-463 Plesiosauria HIasmosauridae Mauisaurus haasli c/d/e/h/l 0 229 AU-285 $-$-h MR^69 Plesiosauria indel. indet. indel. c/h/i 0 229 AU-285 $-$-h MR-466 Plesiosauria indet. indet. indel. e/h/i 0 309;229 AU-285 $-$-h MR-468 Plesiosauria indet. indel. indel. i 0 229 AU-285 $-$-h MR-470 Plesiosauria indet. indel. indel. e 0 229 AU-285 $-S-h MR-467 Plesiosauria Polycotylidae indel. indel. c/i 0 229 TABLE G.2 Marine Reptile Database Taxa

Loc Map MMR Order Family Genus Species Complete . Rererence ID Interval ID

AU-320 $-$-h MR-541 Squamala Mosasauridae indel. indel. i 5 159 AU-41 $-h-g MR-73 Ichthyosauria Platypterygidae Plalyplerygius longnwni b/c/d/e/f/h/i 6/4 305 AIMI $-h-g MR-1484 Plesiosauria Pliosauridae Kronosaurus queenslandicus b/c/d 6 196 AlJ-450 j-j-h MR-707 Plesiosauria l:lasmosauridac Mauisaurus haasli h/i 5 310 AU-451 j-j-h MR-714 PlesiosBuria Elasmosauridae indel. indel. e/i 5/6 310 AU-451 j-j-h MR-708 Plesiosauria l:lasmosauridae Mauisaurus haasli dh 5/6 310 AU-451 J-j-h MR-720 Plesiosauria indel. indel. indel. dhli 5 310 AU-451 j-j-h MR-717 Plesiosauria Polycoiylidae indel. indel. dm 6 310 AU-451 j-j-h MR-729 Squaniata Mosasauridae indel. indel. d 2 310 AU-451 j-j-h MR-724 Squamala Mosasauridae Prognalhodon waiparaensis b/c/d/e/f 6 310 AU-452 j-j-h MR-715 Plesiosauria l-lasmosauridae indel. indel. e/h/i 6 310 AU-452 j-j-h MR-709 Plesiosauria Glasmosauridae Mauisaurus haasli h.'i 5 310 AU-452 j-j-h MR-721 Plesiosauria indel. indel. indel. dflhli 5 310 AU-452 j-j-h MR-719 Plesiosauria Polycoiylidae indel. indel. d/e/i 5 310 AU-452 j-j-h MR-728 Squamala Mosasauridae indel. indel. b/d/e/f/z 6 310 AU-452 j-j-h MR-726 Squamala Mosasauridae Mosasaurus mokoroa 1 6 310 AU-452 j-j-h MR-723 Squamala Mosasauridae Tanivvhasaurus oweni b/c/d 6 310 AU-452 j-j-h MR-722 Squamala Mosasauridae Tylosaurus haumuriensis b/c/d 6 310 AU-453 j-j-h MR-710 Plesiosauria Elasmosauridae Mauisaurus haasli h/i 5/6 310 AU-454 j-j-h MR-716 Plesiosauria Elasmosauridae indel. indel. c/f/g 6 310 AU-454 j-j-h MR-917 Plesiosauria Elasmosauridae indel. indel. c/h 0 309 AU-454 j-j-h MR-7II Plesiosauria Elasmosauridae Mauisaurus haasli e/h/i 6 310 AU-454 j-j-h MR-918 Plesiosauria indel. indel. indel. e 0 309 AU-454 j-j-h MR-718 Plesiosauria Polycoiylidae indel. indel. i 5 310 AU-454 j-j-h MR-727 Squamala Mosasauridae indel. indel. c/d 5 310 AU-455 j-j-h MR-712 Plesiosauria Elasmosauridae Mauisaurus haasli i 6 310 AU-456 j-j-h MR-713 Plesiosauria Elasmosauridae Mauisaurus haasli g 6 310 AU-457 j-j-h MR-725 Squamala Mosasauridae Mosasaurus niokuroa hiddle 6 310 AU-509 $-i-h MR-849 Plesiosauria Elasmosauridae alT. keyesi e/f/g/h 6 317 I'uarangisaurus AU-509 $-i-h MR-848 Plesiosauria Elasmosauridae air. keyesi e/h 6 317 Tuarangisaurus AU-509 S-i-h MR-847 Plesiosauria Elasmosauridae cf. Tuarangisaurus keyesi ddldfMi 6 317 AU-509 S-i-h MH-851 Plesiosauria Elasmosauridae indel. indei. d/e/r/g/h/i 5 317 AU-509 $-i-h MR-850 Plesiosauria Elasmosauridae Mauisaurus haasli e/f/h/i 6 317 AU-509 $-i-h MR-846 Plesiosauria Elasmosauridae Tuarangisaurus keyesi b/c/d/e 6 317 AU-509 $-l-h MR-852 Plesiosauria Pliosauridae indel. indel. d/e/f/h/i 5 317 AU-509 $-l-h MR-839 Squamala Mosasauridae indel. indel. h 5/6 316 OJ KJ -1^ TABLE G.2 Marine Reptile Database Taxa

Loc Map MMR Preser­ Order Family Genus Species Compleie Referenc ID Interval ID vation

AU-509 $-i-h MR-838 Squamala Mosasauridae Mosasaurus alT. mangahouangac dd/df 5/6 316;20 AU-509 $-i-h MR-837 Squamala Mosasauridae Mosasaurus flemingi b/c/d/e/r 5/6 316 AU-509 $-l-h MR-853 Squamala Mosasauridae Mosasaurus mangahouangac hldd/dfli 6 315;20 AU-509 $-i-h MR-840 Squamala Mosasauridae Prognalhodon aff. overtoni b/c/d 6 316 AU-509 $-i-h MR-836 Squamala Mosasauridae Rikisaurus lehoensis b/c/d/e 6 316 AU-51 S-h-g MR-III Ichihyosauria Plaiyplerygidae Plaiyplerygius ausiralis b/d/e/i 6 17 AU-635 $-$-h MR-1139 Plesiosauria indei. indel. indel. a 7 98 AU-635 $-$-h MR-I49I Pleslosauria indel. indel. indel. a 7 98 AU-635 $-$-h MR-1492 Squamala Mosasauridae indel. indel. e 0 99 AU-68 $-a-c MR-135 Ichihyosauria indel. indel. indel. c/r/z 4/5/6 97 AU-69 $-a-c MR-136 Ichihyosauria Shaslasauridae indel. indel. c/d 3 (55; 48 AU-70 $-h-g MR-137 Ichihyosauria indel. indel. indel. di 4/5 97 AU-707 b-b-c MR-1457 Ichihyosauria indel. indel. indel. d/c/r 4 170 AU-707 b-b-c MR-1354 Ichihyosauria Shonisauridae Shonisaurus sp. c/d 3 170;48 AU-707 b-b-c MR-1300 Plesiosauria Plesiosauridae indel. indel. b/c/d 4 54 AU-740 $-a-c MR-1356 Ichihyosauria indel. indel. indel. e 0 48; 99 AU-753 $-a-b MR-1382 Ichihyosauria indel. indel. indel. % 0 48 AU-825 h-h-h MR-1487 Ichihyosauria indel. indel. indel. b/c/d 0 99 AU-826 $-$-b MR-1488 Ichihyosauria indel. indel. indel. d/e 6 99 AU-827 j-j-h MR-1489 Plesiosauria Llasmosauridae indel. indel. e 0 99 AU-827 j-j-h MR-1490 Squamala Mosasauridae indel. indel. d 0 99 •AU-828 S-S-d MR-1493 Plesiosauria Pliosauridae afl'. Lepiocleidus indel. e/d/i 5 31; 196 AU-829 $-h-g MR-1495 Ichihyosauria Plaiyplerygidae Plaiyplerygius indel. e/h/2 4 201 AU-829 $-h-g MR-1494 Plesiosauria Elasmosauridae indel. indel. e/h/i/z 4 201 AU-84 $-$-<1 MR-163 Plesiosauria Plesiosauridae indel. indel. e 3 295 AU-856 8-S-8 MR-1537 Ichihyosauria indel. indel. indel. e 0 197 AU-856 g-g-g MR-1536 Plesiosauria indel. indel. indel. z 0 197 AU-857 g-g-g MR-1539 Ichihyosauria indel. indel. indel. d/e 0 197 AU-857 8-B-B MR-IS38 Plesiosauria indel. indel. indel. $ 0 197 AU-97 6-S-B MR-1485 Ichihyosauria indel. indel. indel. S 0 196 AU-97 g-g-g MR-180 Plesiosauria indel. indel. indel. d 2 226 AU-97 8-8-g MR-182 Plesiosauria Plesiosauroidea indel. indel. d/e/f/h/i 3 226; 309 AU-97 g-g-g MR-179 Plesiosauria Polycolylidae Dolichorhynchops sp. d/e/i/z 2 226;196 AU-98 B-g-g MR-I8I Plesiosauria Polycoiylidae indel. indel. c/z 2 226 AU-99 $-h-g MR-188 Plesiosauria Hlasmosauridae indel. indel. e 4 226 AU-99 $-h-g MR-183 Plesiosauria indel. indel. indel. e 2 226; 309 EU-106 $-d-e MR-192 Plesiosauria Plesiosauridae Plesiosaurus sp. e/i 5 312 l-U-107 h-h-h MR-193 Plesiosauria Llasmosauridae indel. indel. e 0 229 U) K) TABLE G.2 Marine Reptile Database Taxa

Loc Map MMR Order Family Genus Spccies Coniplcle . ' Rererence ID nierval ID

KU 108 $-i-h MR-194 I'lesiusauria indei. indet. indet. e 0 229 UU 109 $-h-g MR-I9S Plesiosauria I'.lasmosauridae indet. indet. d/c/h/i 0 309; 229 EU no $-h-g MR-893 Ichlhyosauria Plaiyplerygidae Plalyplerygius "canipylodon" c/d S 82; 173 EU no $-h-g MR-201 Plesiosauria indei. indet. indet. e 0 309; 229; 34 EU no S-h-g MR-196 Plesiosauria Pliosauridae indet. indet. e 0 309; 229 EU no $-h-g MR-199 Plesiosauria Pliosauridae Polyptychodon inlerruptus b/e/i 0 229 EU in 8-g-g MR-197 Plesiosauria Elasmosauridae indet. indet. c/h 0 309;229 EU 111 g-g-g MR-200 Plesiosauria Pliosauroidea indet. indet. d/c/i 0 309; 229 EU 112 r-r-g MR-198 Plesiosauria Pliosauroidea indet. indet. e 0 309; 229; 34 EU 113 $-$-g MR-202 Plesiosauria Pliosauroidea indet. indet. e 0 309; 229 *EU 114 $-r-g MR-203 Plesiosauria Pliosauridae Leptocleidus supersles a 7 229; 34 EU lis e-e-f MR-88S Ichlhyosauria indei. indet. indet. e S 82; 176 EU 115 e-e-f MR-889 Ichthyosauria indei. indet. indet. f s 82 EU IIS e-e-f MR-207 Ichlhyosauria indei. indet. indet. e 0 176; 34 EU lis e-e-f MR-205 Plesiosauria Ciyploclididae Colymbosaurus? truchanterius e/l 0 229; 40; 34 EU 115 e-e-f MR-781 Plesiosauria Cryploclididae Cryptocleidus aff. richardsoni S 0 81;40 EU lis e-e-f MR-204 Plesiosauria indei. indet. indet. e 0 229; 40 EU lis e-e-f MR-788 Plesiosauria indet. indet. indet. d/z 0 81 EU lis e-e-f MR-790 Plesiosauria indei. indei. indet. z/h 0 81 EU lis e-e-f MR-206 Plesiosauria Pliosauridae ?Pliosaurus brachydeirus h/i 0 34 EU lis e-c-f MR-794 Plesiosauria Pliosauridae Pliosaurus brachydeirus h/i 0 81 EU lis e-c-f MR-Sll Plesiosauria Pliosauridae Pliosaurus sp. h 0 81 EU 116 e-e-f MR-208 Plesiosauria indet. indei. indet. e 0 229; 40 EU 117 $-c-r MR-892 Ichlhyosauria Ichihyosauridae Brachyplerygius extremus i 5/6 82 EU 117 $-e-r MR-873 Ichlhyosauria indei. indet. indet. e 5 82; 176 EU 117 S-e-f MR-907 Ichlhyosauria indet. indet. indet. b/e/h 5 82 EU 117 $-c-r MR-888 Ichlhyosauria indei. indet. indet. c/d/e/h 5 82 EU 117 $-e-f MR-883 Ichlhyosauria indet. indet. indet. e 5 82; 176 EU 117 S-e-r MR-786 Plesiosauria indet. indet. indet. d 0 81 EU 117 $-c-f MR-770 Plesiosauria indet. indet. indet. e 0 81;40 EU 117 MR-209 Plesiosauria indet. indet. indet. e 0 229; 40 EU 117 s-e-f MR-235 Plesiosauria Pliosauridae indet. indet. e 0 229 EU 117 $-c-f MR-802 Plesiosauria Pliosauridae I.iopleurodun '.'macromerus d 0 8I;291 EU 117 S-e-f MR-791 Plesiosauria Pliosauridae Pliosaurus brachydeirus d/e 0 81 EU 117 S-e-f MR-809 Plesiosauria Pliosauridae Pliosaurus sp. S 0 81 EU 117 S-e-f MR-813 Plesiosauria Pliosauridae Pliosaurus sp. e 0 81 EU 118 S-e-f MR-16n Ichlhyosauria Ichihyosauridae Brachyplerygius extremus b/c/d/e/f/h 6/7 187 EU 118 S-e-f MR-908 Ichlhyosauria indet. indet. indet. b/c/d/z 5 82 u> to 0\ TABLE G.2 Marine Reptile Database Taxa

Loc Map MMR Order l-amily Genus Species Complete . ' Rcrcrence ID nlcrval ID * ' vntmn l-U-118 $-e-f MR-874 Ichthyosauria indet. indet. indet. e 5 82; 176 BU-II8 $-e-f MR-879 Ichlhyosauria indet. indet. indet. e 6 82;176 HIJ-118 $-c-r MR-904 Ichthyosauria Platyptcrygidac Nannopterygius enthekiodon c/d/c/r/h/i 5 82 EU-II8 $-c-r MR-777 Plesiosauria Cryptoclididae ".'Colymbosaurus trochanterius s 0 81 I:U-II8 $-c-f MR-211 Plesiosauria Cryptoclididae Colymbosaurus Uochanlerius erti/i 0 229; 40 EU-II8 $-c-r MR-784 Plesiosauria indet. indet. indet. d 0 81 EU-II8 s-c-r MR-771 Plesiosauria indet. indet. indet. c 0 8I;40 EU-II8 $-c-f MR-210 Plesiosaurii indet. indet. indet. e/r/i 0 81; 229; 40 EU-II8 S-c-f MR-799 Plesiosauria Pliosauridae I.iopleurodon macromerus d 0 81; 291 EU-II8 $-e-f MR-797 Plesiosauria Pliosauridae l.iopleurodon macromerus h/i 0 81; 291 EU-II8 $-c-f MR-796 Plesiosauria Pliosauridae Pliosaurus brachyspondylus e 0 81 EU-II8 S-e-f MR-806 Plesiosauria Pliosauridae Pliosaurus sp. S 0 81 EU-II8 $-«-r MR-812 Plesiosauria Pliosauridae Pliosaurus sp. c 0 81 EU-II9 $-c-r MR-212 Plesiosauria Cryptoclididae Col>7nbosaurus trochanterius e 0 229; 40 EU-II9 s-c-r MR-228 Plesiosauria Pliosauridae Pliosaurus brachyspondylus c/e/h/i 0 229 EU-12 $-8-b MR-IS Ichlhyosauria Shastasauridae Shastasaurus neubigi b/c/d/e/r/g/h 6 254 /i EU-120 S-e-f MR-213 Plesiosauria Cryptoclididac Colymbosaurus? trochanterius e/l 0 229 EU-I2I $-e-f MR-872 Ichthyosauria indet. indet. indet. e 5 82;176 EU-I2I S-c-r MR-881 Ichthyosauria indet. indet. indet. c 5 82; 176 EU-121 S-c-r MR-214 Plesiosauria indet. indet. indet. e 0 229; 40 EU-122 S-c-r MR-242 Plesiosauria indet. indet. indet. e 0 229 EU-122 S-e-r MR-215 Plesiosauria indet. indet. indet. c 0 229; 40 EU-122 s-c-r MR-239 Plesiosauria indet. indet. indet. c 0 229;40 EU-122 s-c-r MR-238 Plesiosauria Pliosauridae indet. indel. h 0 229 EU-122 s-c-r MR-236 Plesiosauria Pliosauridae indet. indet. i 0 229 EU-122 s-c-r MR-229 Plesiosauria Pliosauridae l.iopleurodon macromerus c/e/h/i 0 229 EU-122 s-e-r MR-814 Plesiosauria Pliosauridae Pliosaurus sp. c 0 81 EU-123 S-d-c MR-216 Plesiosauria Cryptoclididae Cryptoclidus curymerus e/h/i 0 229;40 EU-123 S-d-c MR-222 Plesiosauria Elasmosauridae Muraenosaurus Moclis a 0 229; 40 EU-123 S-d-c MR-220 Plesiosauria Elasniosauridae Muraenosaurus lecdsii a 0 229 EU-123 S-d-e MR-223 Plesiosauria Elasmosauridae Muraenosaurus Icedsii a 0 229; 40 EU-123 S-d-c MR-224 Plesiosauria Elasmosauridae Muraenosaurus lecdsii a 0 229; 40 EU-123 S-d-c MR-225 Plesiosauria Elasmosauridae Tricleidus seclcyi a 0 229; 40 EU-123 S-d-c MR-230 Plesiosauria Pliosauridae l.iopleurodon ferox a 0 229 EU-123 S-d-e MR-231 Plesiosauria Pliosauridae l.iopleurodon pachydcirus a 0 229 EU-123 S-d-c MR-232 Plesiosauria Pliosauridae Peloneustcs philarchus a 0 229 EU-123 S-d-c MR-233 Plesiosauria Pliosauridae Pliosaurus andrewsi a 0 229 U) NJ TABLE G.2 Marine Reptile Database Taxa

Loc Map MMR „ . II-. I.I ir. Order romily Genus Species Complete . " Reference ID Interval ID ' ' vnlinn

I-IJ.123 S-d-e MR-240 I'lesiosauria Pliosauridae Simolcstcs vorax a 0 229 EU-124 $-d-e MR-217 Plcslosauria Cryptoclididae Cryptoclidus eur>inerus i 0 229 l-U-125 S-d-c MR-218 Plesiosauria Cryptoclididae Cryptoclidus eurymerus a 0 229; 40 I:U'I26 S-d-c MR-778 PIcsiosauria Cryptoclididae Cryptoclcidus cuiymerus e 0 81:40 KU.I26 $-d-« MR-779 Plesiosauria Cryptoclididae Cryptoclcidus richardsoni S 0 8I;40 EU-126 $-d-« MR-219 Plesiosauria Cryptoclididae Cryptoclidus richardsoni e/h 0 229 EU-126 $-d-e MR-769 Plesiosauria Elasmosauridae Muraenosauius lecdsii c 0 8I;40 EU'I26 $-d-e MR-234 Plesiosauria Pliosauridae indet. indet. c 0 81,229 EU-126 $-d-c MR-805 Plesiosauria Pliosauridae Liopleurodon Vferoji i 0 81; 291 EU-126 S-d-e MR-804 Plesiosauria Pliosauridae l.iopleurodon ferox d 0 81; 291 EU-127 $-d-c MR-221 Plesiosauria Elasmosauridae Muraenosaurus lecdsii c 0 229; 40 EU-128 S-d-c MR-226 PIcsiosauria Plcsiosauridae indet. indet. b/d/cAi/i 0 229 EU-129 S-c-f MR-227 PIcsiosauria Pliosauridae Pliosaurus brachydcirus c/c/i 0 229 EU-130 S-c-f MR-880 Ichthyosauria indet. indet. indet. c 5 82; 176 EU-130 S-c-f MR-871 Ichthyosauria indet. indet. indet. c 5 82; 176 EU-130 S-c-f MR-878 Ichthyosauria indet. indet. indet. b/c/z 6 82; 176 EU-130 S-c-f MR-237 Plesiosauria Pliosauridae indet. indet. i 0 229 EU-13« $-e-f MR-241 Plesiosauria indet. indet. indet. i 0 229 EU-I3I S-c-f MR-795 Plesiosauria Pliosauridae Pliosaurus brachyspondylus e 0 81 EU.I32 S-c-d MR-1223 Ichthyosauria indet. indet. indet. S 0 35 EU-132 S-c-d MR-1222 Ichthyosauria Leptopterygidae Eurhinosaurus longirostris a 7 35; 179 EU-132 S-c-d MR-I22I Ichthyosauria Tcmnodontosaurid Tcmnodontosaurus burgundiae q 6/7 177; 35; 186 &c EU-132 S-c-d MR-243 Plesiosauria Elasmosauridae Microcleidus homalospondylus a 0 229 EU.I32 S-c-d MR-254 Plesiosauria indet. indet. indet. a 0 229 EU-133 S-c-d MR-244 Plesiosauria Elasmosauridae Microcleidus macroptcrus a 0 229 EU-133 S-c-d MR-255 PIcsiosauria Pliosauridae Rhomaleosaurus /etiandicus a 0 229; 73 EU.I34 S-c-d MR-864 Ichthyosauria Leptoptcrygiidae Leptonectes solei a 7 181;186 EU-134 S-c-d MR-245 PIcsiosauria Plcsiosauridae Attenborosaurus conybeari a 0 229; 15 EU-135 S-c-d MR-246 Plesiosauria indet. indet. indet. e 0 229 EU-136 S-c-d MR-868 Ichthyosauria Ichthyosauridae Ichthyosaurus breviceps a 7 82; 174 EU-136 S-c-d MR-866 Ichthyosauria Ichthyosauridae Ichthyosaurus communis a/q 6/7 82; 174; 34 EU-136 S-c-d MR-867 Ichthyosauria Ichthyosauridae Ichthyosaurus communis b/c/d/c/h/i 6 82; 174 EU-136 S-c-d MR-869 Ichthyosauria Ichthyosauridae Ichthyosaurus conybeari b/c/d/e 6 82; 174 EU-136 S-c-d MR-903 Ichthyosauria indet. indet. indet. h 5 82; 179 EU-136 S-c-d MR-905 Ichthyosauria indet. indet. indet. e 5 82 EU-136 S-c-d MR-896 Ichthyosauria Leptopterygidae Leptonectes '.'tenuirostris c^ 6 82;186 EU-136 S-c-d MR-900 Ichthyosauria Leptopterygidae Leptonectes sp h 5/6 82; 186 U>

00 TABLE G.2 Marine Reptile Database Taxa

Luc Map MMR Order Family Genus Species Complete . ' Reference ID Interval ID

1:IJ-I36 $-c-d MR-895 Ichlhyosauria l.eploplerygidae Leptoncctes tenuirosiris d/a/b/i SIbn 82; 186 EU-136 $-c-d MR-899 Ichlhyosauria Tcmnodontosaurid Tcmnodontosaurus ?plaiyodon h/z 516 82;186 ae l:U-136 $-c-d MR-902 Ichlhyosauria Tcmnodontosaurid Temnodontosaurus '.'playiodon h 5 82; 175 ae nU-136 $-c-d MR-897 Ichlhyosauria Tcmnodontosaurid rcmnodontosaufus plalyodon c/d/c/i 6 82; 186

UU-136 $-c-d MR-901 Ichlhyosauria Tcmnodontosaurid Tcmnodoniosaurus playtodon q 6 82; 175 ae EU-136 $-c-d MR-265 Plesiosauria indcl. "PIcsiosaurus" macrocephalus a 0 229 EU-136 $-c-d MR-251 PlesiosBuria indet. indcl. indel. e 0 229 EU-136 $-c-d MR-249 Plesiosauria indel. indcl. indel. e/l 0 229 EU-136 $-c-d MR-248 Plesiosauria Plesiosauridae PIcsiosaurus dolichodeirus a 0 229 EU-137 $-b-c MR-250 Plesiosauria indel. indcl. indet. c 0 229; 21 EU-138 $-c-d MR-252 Plesiosauria Pliosauridae Rhomaleosaurus zctlandicus a 0 229; 73 EU-139 $-S-d MR-253 Plesiosauria Pliosauridae Rhomalcosaurus thomtoni a 0 229 EU-140 $-c-d MR-2S6 Plesiosauria incertae sedis Sihcnarosaurus dawkinsi c/h/i 0 229 EU-141 $-e-d MR-258 Plesiosauria indel. "PIcsiosaurus" propinquus a 0 229 EU-141 $-c-d MR-271 Plesiosauria indel. indel. indcl. e 0 229 EU-141 $-c-d MR-257 Plesiosauria indel. Macroplata longirostris b 0 229 EU-142 $-c-d MR-259 Plesiosauria Elasmosauridac Erelmosaurus rugosus c/h/i 0 229 EU-143 $-c-d MR-260 Plesiosauria indel. "PIcsiosaurus" macriKcphalus a 0 229 EU-144 $-c-d MR-898 Ichlhyosauria 1'emnodontosaurid Tcmnodontosaurus plalyodon e 5/6 82 ae EU-144 $-c-d MR-766 Plesiosauria Elasmosauridac '.'Erelmosaurus sp. c 6 81 EU-144 $-c-d MR-762 Plesiosauria Plesiosauridae PIcsiosaurus dolichodcirus $ 0 81 EU-144 $-c-d MR-261 Plesiosauria Pliosauridae Archaeonccirus rostralus a 0 229 EU-145 $-c-d MR-266 Plesiosauria incertae sedis Thalassiodracon hawkinsi hldh 0 229;286 EU-145 $-c-d MR-764 Plesiosauria indel. indel. indet. c/f/li/i 6 81 EU.145 $-c-d MR-262 Plesiosauria Pliosauridae Eurycleidus mcgacephalus a 0 229 EU-146 $-c-d MR-263 Plesiosauria Pliosauridae Macroplata lenuiccps a 0 229 EU-147 $-e-d MR-264 Plesiosauria Pliosauridae Eurycleidus arcualus c/e/h/i 0 81; 229 EU-148 $-b-c MR-1604 Ichlhyosauria indcl. "Ichthyosaurus" sp. c/d/e/i 4 34 EU-148 $-b-c MR-267 Plesiosauria Elasmosauridae Erelmosaurus rugosus e 0 229 EU-148 $-b-c MR-269 Plesiosauria incertae sedis Thalassiodracon hawkinsi c 0 229;286 EU-148 $-b-c MR-268 Plesiosauria indel. indet. indel. d/e 0 229 EU-148 $-b-c MR-270 Plesiosauria indel. indet. indel. d/e 0 229 EU-149 $-d-c MR-272 Plesiosauria indet. indet. indet. c/c/1 0 229 U) to TABLE G.2 Marine Reptile Database Taxa

Loc Map MMR Order Family Genus Specics Complete . Reference ID Interval ID ' vation

EU-150 $-$-h MR-273 Plcsio&auria Elasinosauridac indct. indct. e/i 0 229 EU-I5I $-$-h MR-274 Plesiosauria Elasmosauridac indct. indct. c 0 229 1:U-I52 $-$-h MR-275 PIcsiosauria Polycolylidae indct. indct. c 0 229 EU-153 $-h-B MR-276 Plesiosauria Elasmosauridac indct. indct. c 0 229 EU-153 $-h-g MR-2g4 PIcsiosauria indct. indct. indct. c 0 229 EU-153 $-h-g MR-285 Plesiosauria indel. indct. indet. c 0 229 EU.154 $-h-g MR-277 Plesiosauria Pliosauridac indct. indet. c 0 309,229 EU-154 $-h-g MR-2g3 Plesiosauria Pliosauridac Polyptychodon intemiptus d 0 229 Ell-155 $-f-B MR-278 Plesiosauria Elasmosauridac Brancasaurus brancai a 0 229 EU-155 $-f-g MR-280 Plesiosauria Plcsiosauridac indel. indel. c 0 229 EU-155 $-r-g MR-279 PIcsiosauria Pliosauroidca indel. indel. e 0 309;229 EU-156 $-f-g MR-281 Plesiosauria Pliosauroidea indct. indet. c/i 0 309; 229 Ell-157 $-r-B MR-282 Plesiosauria UlBsmosauridBc? indet. indet. e 0 309;229 EU-159 c-c-f MR-287 Plesiosauria indct. indct. indel. e 0 229;40 EU-159 e-e-f MR-297 Plesiosauria Pliosauridac Pliosaurus brachydcirus e 0 229 r:U-l60 e-c-f MR-288 Plesiosauria Cryptoclididac Colymbosaurus? cf C. trochantcrius e 0 229 EU-I6I $-c-f MR-305 Plesiosauria indct. indct. indel. e 0 229 EU-161 $-c-f MR-304 Plesiosauria indel. indel. indel. e 0 229;40 EU-161 $-e-r MR-289 Plesiosauria indct. indel. indel. e 0 229 EU-161 $-c-l MR-290 Plesiosauria indct. indel. indel. c 0 229; 40 EU-161 $-c-f MR-296 Plesiosauria Plcsiosauridac indel. indet. c 0 229 EU-161 $-«-f MR-303 Plesiosauria Pliosauridac indel. indct. i 0 229 EU-162 $-c-f MR-291 PIcsiosauria indct. indel. indet. c 0 229; 40 EU-162 $-c-r MR-1204 Plesiosauria Pliosauridac Pliosaurus brachyspondylus c 5 25 BU-164 d-d-c MR-293 Plesiosauria indel. indct. indct. b 0 229 EU-164 d-d-c MR-300 Plesiosauria Pliosauridac Lioplcurodon d". L. Tcrox d 0 229 EU-165 $-d-c MR-294 PIcsiosauria Elasmosauridac Muracnosaurus? sp. e 0 229 EU-167 $-c-r MR-298 Plesiosauria Pliosauridac Lioplcurodon macromcrus c 0 229 EU-168 $-«-r MR-299 PIcsiosauria Pliosauridac Lioplcurodon Icrox d 0 229 EU-169 d-d-e MR-301 PIcsiosauria Pliosauridac Pliosaurus andrcwsi d 0 291;229 EU-170 $-d-e MR-302 PIcsiosauria Pliosauridac Liopleurivlon pachydcirus d 0 229 EU-171 c-c-d MR-306 Plesiosauria Elasmosauridac Micrucleidus cf. M. c 0 229 homalospondylus EU-I7I c-c-d MR-310 Plesiosauria incertae scdis Sihenarusaurus dawkinsi h 0 229 EU-171 c-c-d MR-309 Plesiosauria Plcsiosauridac PIcsiosaurus cf. P. dolichodcirus e 0 229 EU-172 $-c-d MR-307 PIcsiosauria Elasmosauridac indet. indct. cyf/g/h/i 0 229;117 EU-173 $-$-d MR-1458 Ichthyosauria indct. indct. indel. b/c/d/z 5 171 EU-173 $-$-d MR-312 Plesiosauria indel. indct. indct. e 0 229 u> u> o TABLE G.2 Marine Reptile Database Taxa

LM Map Order Family Genus Species Complete Pfcscr Rgfefencc IP interval iP ' \ \ vation

EU-173 S-S-d MR-3M Plesiosauria indel. indet. indet. c 0 229 EU.I74 $-b-c MR-IISI Ichihyosauria indet. indel. indel. e 5 21 EU-174 $-b-c MR-313 Plesiosauria indet. indet. indel. e 0 229; 21 EU-174 $-b-c MR-314 Plesiosauria indel. indel. indel. e 0 229;21 EU-175 $-$-b MR-1I4I Ichihyosauria Mixosauridae Mixosaurus sp. e 5 21 EU-175 $-$-b MR-J142 Ichthyosauria Shasiasauridae Cyinbospondylus sp. e 5 21 EU-175 $-$-b MR-316 Plesiosauria indel. indel. indel. e 0 229; 21 EU-176 h-h-h MR-317 Plesiosauria indel. indel. indel. e 0 229 EU'I77 $-c-d MR-318 Plesiosauria Plesiosauridae Plesiosaurus sp. b 0 229 EU-178 $-i-h MR-319 Plesiosauria Pliosauridae Polyptychodon? cf. P. inlerruptus c/d 0 229 EU-179 $-i-h MR-320 Plesiosauria Pliosauridae Polyptychodon cf. P. inlerruptus b 0 229 EU-ISO $-i-h MR-321 Plesiosauria indet. indet. indel. i 0 229 EU-I8I $-i-h MR-322 Plesiosauria indet. indet. indet. c 0 229 EU.I82 $-i-h MR-324 Plesiosauria indel. indet. indet. e 0 229 EU-182 S-i-h MR-323 Plesiosauria indel. indel. indel. e/h 0 229 EU-183 $-i-h MR-325 Plesiosauria indel. indel. indel. c 0 229 EU-184 $-$-g MR-326 Plesiosauria Elasmosauridae indel. indel. e 0 229 EU-185 g-g-g MR-328 Plesiosauria Plesiosauridae indet. indel. e 0 229 EU-185 g-g-g MR-327 Plesiosauria Plesiosauridae indel. indel. c 0 229 EU-186 $-$g MR-329 Plesiosauria Pliosauridae Polyptychodon cf. P. inlerruptus d 0 229 EU-187 $-$-g MR-330 Plesiosauria indet. indet. indet. e 0 229 EU-189 $-«-f MR-332 Plesiosauria Pliosauridae Liopleurodon cf 1.. ferox d 0 229 EU-190 d-d-e MR-333 Plesiosauria Pliosauridae l.iopleurodon ferox b/e/i 0 229 EU.19I $-«-r MR-334 Plesiosauria indel. indet. indet. d/e/i 0 229 EU-192 c-c-f MR-335 Plesiosauria indet. indet. indel. e 0 229 EU-193 $-c-d MR-336 Plesiosauria Plesiosauridae Plesiosaurus guilelmi impcratoris a 0 229;285 EU-193 $-c-d MR-337 Plesiosauria Plesiosauridae Plesiosaurus guilelmi impcratoris a 0 229 EU-193 $-c-d MR-344 Plesiosauria Pliosauridae Rhomaleosaurus victor a 0 229 EU-195 $-e-d MR-339 Plesiosauria Plesiosauridae Plesiosaurus dolichodcirus e^ 0 229 EU-196 $-c-d MR-340 Plesiosauria Plesiosauridae Plesiosaurus cf P. dolichodcirus i 0 229 EU-197 $-c-d MR-341 Plesiosauria indet. indel. indet. e/i 0 229 EU-198 $-$-c MR-342 Plesiosauria indel. indet. indet. c 0 229 EU-199 $-$-b MR-I36I Ichihyosauria indet. indel. indet. $ 0 48 EU-199 $-$-b MR-1363 Ichthyosauria Mixosauridae Mixosaurus sp S 0 48 EU-199 $-$-b MR-1360 Ichthyosauria Shasiasauridae Cyinbospondylus sp. $ 0 48 EU-199 $-$-b MR-343 Plesiosauria indel. indet. indet. e 0 229 EU-199 $-$-b MR-349 Plesiosauria Pistosauridae Pislosaurus longaevus b/df/h/i 7 229;288 EU-20 $-a-b MR-1387 Ichthyosauria incertae sedis Pessosaurus sp. e 0 48 U) TABLE G.2 Marine Reptile Database Taxa

I.oc Map MMR Order Family Genus Spccies Complete . Rererence ID 1Interval ID • ' vntinn

EU-20 $-a-b MR-1384 Ichlhyosauria Mixosauridae Mixosaurus atavus S 0 48 EU-20 $-a-b MR-1373 Ichihyosauria Shastasauridae Cymbospondylus sp. e 0 48 l:U-200 $-c-d MR-345 Plesiosauria Pliosauridae Eurycleidus niegacephalus a 0 229 UU-201 $-b-c MR-346 Plesiosauria indet. indet. indet. d/e 0 229 l-lJ-203 $-c-d MR-350 Plesiosauria indet. indet. indet. i 0 229 BU-204 $-$-d MR-351 Plesiosauria indet. indet. indet. e 0 229 l-U-205 $-c-d MR-352 Plesiosauria indet. indet. indet. i 0 229 EU-206 $-i-h MR-353 Plesiosauria Elasmosauridae indet. indet. e 0 309,229 l-U-207 $-i-h MR-356 Plesiosauria Elasmosauridae indet. indet. e 0 309;229 r:U-207 $-i-h MR-354 Plesiosauria Elasmosauridae indet. indet. e 0 309; 229 EU-21 S-a-b MR-1378 Ichlhyosauria indet. indet. indet. S 0 48 EU-21 $-a-b MR-1379 Ichthyosauria Mixosauridae cf. Mixosaurus cf M. atavus S 0 48 EU-21 S-a-b MR-1374 Ichthyosauria Shastasauridae Cymbospondylus sp. b/e/r 4 48 EU-210 $-i-h MR-359 Plesiosauria Pliosauroidea indet. indet. d/e/i 0 309; 229 EU-23 $-a-b MR-1380 Ichlhyosauria indet. indet. indet. 1 0 48 EU-23 $-a-b MR-I38I Ichlhyosauria Mixosauridae Mixosaurus cr. M. atavus S 0 48 EU-23 $-a-b MR-1390 Ichthyosauria Shastasauridae indet. indet. h/1 0 48 EU-244 $-c-f MR-4II Plesiosauria indet. indet. indet. e/h/i 6 227; 229 El)-245 $-$-r MR-412 Plesiosauria Plesiosauridae indet. indet. e 0 229 EU-246 e-e-f MR-413 Plesiosauria Plesiosauridae indet. indet.

Loc Map MMR „ , in. 1.1 11-. Order Family Genus Species Complete .. " Rererencc ID Interval II) ' ' vaiion

ac I:U-300 $-c-d MR-505 Ichlhyosauria Stcnopierygidae Sienopicrygius quadriscissus a 7 177 OU-301 $-c-d MR-506 Ichlhyosauria Stcnopierygidae Sienopicrygius quadriscissus a 7 177 ElJ-302 $-c-d MR-512 Ichlhyosauria Lcpioplcrygidac liurhinosaurus longirosiris b/c 6 177; 184 l:U-302 $-c-d MR-508 Ichlhyosauria Stcnopierygidae Sienopterygius haufTianus b 6 177 EU-302 $-c-d MR-510 Ichlhyosauria Stcnopierygidae .Sienopicrygius longipcs a 7 177 EU-302 S-c-d MR-5II Ichlhyosauria Slenoptcrygidae Sienopicrygius macrophasma a 7 177 EU-302 $-c-d MR-509 Ichlhyosauria Stcnopierygidae Sienopicrygius megalorhinus a/b 6/7 177 EU-302 $-c-d MR-507 Ichlhyosauria Stcnopierygidae Sienopicrygius quadriscissus a 7 177 EU-302 $-c-d MR-513 Ichlhyosauria Tcmnodonlosaurid Tcmnodontosaurus burgundiac b/c/f 6 177.186 ae EU-303 $-c-d MR-514 Ichlhyosauria Stcnopierygidae Sienopterygius magacephalus a 7 177 EU-304 $-c-d MR-515 Ichlhyosauria Stenoplcrygidac Sienopterygius megalorhinus a 7 177 EU-305 $-c-d MR-516 Ichlhyosauria Lcpioplcrygidac Eurhinosaurus longirosiris a 7 177; 184 EU-305 $-c-d MR-517 Ichlhyosauria Tcmnodonlosaurid Tcmnodontosaurus burgundiae a 7 177; 186 ac EU-306 $-c-d MR-518 Ichlhyosauria Tcmnodonlosaurid Tcmnodontosaurus burgundiae b 6 177; 186 ac EU-307 $-c-d MR-519 Ichlhyosauria Slcnoptcrygidae Sienopterygius haufTianus a 7 177 EU-308 $-c-d MR-521 Ichlhyosauria Stcnopierygidae Sienopterygius cunciccps a 7 177 EU-308 $-c-d MR-520 Ichlhyosauria Slenoptcrygidae Sienopterygius haufTianus b/c 6 177 EU-309 $-c-d MR-522 Ichlhyosauria Stenoplcrygidac Sienopterygius megalorhinus a 7 177 EU-310 $-c-d MR-524 Ichlhyosauria Stcnopierygidae Sienopicrygius cunciccps a 7 177 EU-310 $-c-d MR-523 Ichlhyosauria Slenoptcrygidae Sienopterygius quadriscissus a 7 177 EU-3II $-c-d MR-525 Ichlhyosauria Tcmnodonlosaurid Tcmnodontosaurus burgundiac a 7 177; 186 ac EU-312 $-c-d MR-526 Ichlhyosauria Tcmnodonlosaurid Tcmnodonlosaurus burgundiac b 6 177;186 ae EU-36 d-d-c MR-933 Ichlhyosauria Ichihyosauridac Ophihalmosaurus icenicus q 6 4 EU-36 d-d-c MR-940 Plesiosauria Cryploclididae Cryploclidus curymerus q 6 4; 40 EU-36 d-d-c MR-937 Plcsiosauria Elasmosauridac Muraenosaurus bcloclis q 6 4; 40 EU-36 d-d-c MR-934 Plesiosauria Elasmosauridac Muraenosaurus lecdsii q 6 4 EU-36 d-d-c MR-935 Plcsiosauria Elasmosauridac Muraenosaurus lecdsii q 6 4; 40 EU-36 d-d-c MR-936 Plcsiosauria Elasmosauridac Muraenosaurus lecdsii q 6 4; 40 EU-36 d-d-e MR-938 Plcsiosauria Elasmosauridac Muraenosaurus Sp. b/c/r/g/h/i 6 4; 40 EU-36 d-d-c MR-939 Plesiosauria Elasmosauridac Triclcidus seeleyi a 6 4 EU-36 d-d-c MR-61 Plesiosauria Pliosauridae cf. Lioplcurodon sp. b/c/d/e/f/h/i 6 160 EU-36 d-d-c MR-941 Plesiosauria Pliosauridae Lioplcurodon fcrox q 6 5; 291 TABLE G.2 Marine Reptile Database Taxa

Loc Map MMR Order Family Cienus Specics Complele Pfcs*:r- Reference ID Interval ID ' vation

ElJ-36 d-d-e MR-943 Plesiusauria Pliosauridae Peloneustes philarchus M 6 5 F.U-36 d-d-c MR-944 Plesio&auria PUosauridae Pliosaurus andrcNvsi q 6 5.291 EU-36 d-d-c MR-942 Plesiosauria Pliosauridae Simolestes vorax q 6 5 EU-395 h-h-h MR-620 Ichlhyosauria Platypterygidae Platypterygius sp. b/c/d/c/f/i/z 4 29 EU-395 h-h-h MR-1070 Plesiosauria Pliosauridae Polyptychodon sp d 2 22 l-U-397 j-j-h MR-624 Squamata Mosasauridae VMosasaurus sp. d/e 0 212 EU-417 $-c-f MR-657 Plesiosauria Cryptoclididae Colymbosaurus trochanterius e/f/gm/i 7 41 EU-418 $-c-d MR-658 Ichthyosauria Ichthyosauridae Iciithyosaurus communis b/e/r/i 5 327;177 EU-420 $-c-d MR-660 Ichlhyosauria indet. indet. indet. b/c/d 5 327 EU-421 $-c-d MR-661 Ichthyosauria indet. indet. indet. b/d 5 327 EU-422 c-c-d MR-662 Ichthyosauria indet. indet. indet. e 5 327 EU-423 c-c-d MR-663 Ichthyosauria Ichthyosauridae Ichthyosaurus communis e 5 327; 76; 174 EU-424 c-c-d MR-665 Ichthyosauria Ichthyosauridae Ichthyosaurus communis e 0 76; 174 EU-424 c-c-d MR-666 Ichthyosauria Stenopteiy'gidae Stenopterygius megalorhinus d/e/f/i 0 76; 177 EU-425 $-c-d MR-667 Ichthyosauria Stenopterygidae Sienopterygius sp. e 0 76 EU-426 $-d-c MR-668 Ichthyosauria Stenopteiygidae Stenopterygius .'megalorhinus e 0 76; 177 EU^27 c-c-d MR-669 Ichthyosauria Ichthyosauridae Ichthyosaurus communis e 0 76; 174 EU-428 $-c-d MR-670 Ichthyosauria indet. indet. indet. dm 0 70 EU-429 $-c-d MR-67I Ichthyosauria indet. indet. indet. e/r/i 0 70 EU-430 $-c-d MR-672 Ichthyosauria indet. indet. indel. z 0 70 EU-431 $-c-d MR-673 Ichthyosauria indet. indet. indei. z 0 70 EU-432 $-c-d MR-674 Ichthyosauria indet. indet. indet. z 0 70 EU-433 $-c-d MR-675 Ichthyosauria indet. indet. indel. c/f 0 70 EU-434 S-c-d MR-676 Ichthyosauria indet. indet. indet. e/f 0 70 EU-435 $-$-d MR-677 Ichthyosauria indet. indet. indel. 7 84 EU-435 $-$-d MR-682 Ichthyosauria indet. indet. indel. 7 84 EU-435 $-$-d MR-681 Ichthyosauria indet. indet. indel. 7 84 EU-435 $-$-d MR-680 Ichthyosauria indet. indet. indel. 7 84 EU-435 $-$-d MR-678 Ichthyosauria indet. indet. indel. 7 84 EU-435 $-$-d MR-679 Ichthyosauria indet. indet. indel. 7 84 EU-436 $-d-e MR-683 Ichthyosauria Ichthyosauridae indet. indel. h 5 84 EU-437 $-c-f MR-684 Ichthyosauria Ichthyosauridae Opiithalmosaums sp. d/e/r/h/i 84 EU-438 $-e-f MR-685 Ichthyosauria indet. indet. indet. i 7 85 EU-439 $-«-r MR-686 Plesiosauria Cryptoclididae Colymbosaurus trochanterius a 7 292 EU-439 $-c-f MR-687 Plesiosauria Pliosauridae indet. indet. i 5 292 EU-440 S-e-f MR-703 Ichthyosauria indet. indet. indet. e/f 5 176; 292 EU-440 $-c-f MR-702 Ichthyosauria indet. indet. indet. e/h 7 292 EU-440 $-e-f MR-688 Plesiosauria Cryptoclididae Colymbosaurus sp. e 5 292 u> u> TABLE G.2 Marine Reptile Database Taxa

Loc Map MMR Prcscf" Order Family Genus Species Complete .. Reference ID 1Intenal ID _ ' valion l-U-440 $-e-f MR-689 Plesiosauria Pliosauridae indet. indet. i 5 292 EU-441 S-e-f MR-692 Ichlhyosauria indet. indet. indet. d/e/f 5 292 EU-441 $-c-f MR-690 Plesiosauria Ciyptoclididae Colymbosaurus sp. b/z 5 292;42 l-U-441 S-e-f MR-691 Plesiosauria indet. indet. indet. d/e/f 5 292 EU-442 s-e-r MR-695 Plesiosauria indet. indet. indet. f 5 292 EU-442 s-c-r MR-694 Plesiosauria Pliosauridae Pliosaurus '.'brachyspondylus e 6 292 EU-443 s-e-r MR-696 Plesiosauria Pliosauridae indet. indet. d 5 292 EU-444 $-c-r MR-697 Plesiosauria indet. indet. indet. e 5 292 EU-445 $-c-f MR-700 Ichlhyosauria indet. indet. indet. b 5 292 EU-445 S-c-f MR-699 Plesiosauria indet. indet. indet. e 5 292 EU-446 $-e-r MR-701 Ichlhyosauria indet. indet. indet. h/i 6 292 EU-448 s-e-f MR-705 Plesiosauria Pliosauridae Peloneustes sp. b/c/e/f/h/L'z 6 32 EU-449 $-s-r MR-706 Ichlhyosauria indet. indet. indet. e/lVi 6 83 EU-458 j-j-h MR-730 Squamata Mosasauridae Mosasaurus holTmanni S 0 237 EU-459 j-j-h MR-731 Squamata Mosasauridoe Mosasaurus holTmanni S 0 237 EU-46 h-h-h MR-106 Ichlhyosauria Platypterygidae Platypterygius "campylodon" b/c 6 17 EU^60 j-j-h MR-732 Squamata Mosasauridae Mosasaurus holTmanni S 0 237 EU-461 j-j-h MR-733 Squamata Mosasauridae Mosasaurus hofTmanni S 0 237 EU-462 j-j-h MR-734 Squamata Mosasauridae Mosasaurus conodon s 0 237 EU-462 j-j-h MR-741 Squamata Mosasauridae Plioplatecarpus houzeaui s 0 237 EU-462 j-j-h MR-742 Squamata Mosasauridae Prognathodon solvayi s 0 237 EU-463 j-j-h MR-739 Squamata Mosasauridae Hainosaurus bemardi s 0 237 EU-463 j-j-h MR-735 Squamata Mosasauridae Mosasaurus conodon $ 0 237 EU-463 j-j-h MR-743 Squamata Mosasauridae Prognathodon solvayi s 0 237 EU-464 j-j-h MR-740 Squamata Mosasauridae Hainosaurus bemardi s 0 237 EU-464 j-j-h MR-736 Squamata Mosasauridae Mosasaurus conodon s 0 237 EU-465 j-j-h MR-737 Squamata Mosasauridae Mosasaurus conodon s 0 237 EU-466 j-j-h MR-738 Squamata Mosasauridae Carinodens frassi s 0 298; 237 EU-47 B-g-S MR-107 Ichthyosauria Platypterygidae Platypterygius pladydactylus a 6 17 EU-475 $-e-f MR-887 Ichlhyosauria indet. indet. indet. e 5 82; 176 EU-475 $-«-f MK-906 Ichlhyosauria indet. indet indet. e 5 82 EU-475 S-c-f MR-798 Plesiosauria Pliosauridae l.iopleurodon macromerus c/d 0 81; 291 EU-475 S-e-f MR-808 Plesiosauria Pliosauridae Pliosaurus sp. S 0 81 EU-479 S-S-il MR-765 Plesiosauria indet. indet. indet. e 5 81 EU-48 B-g-E MR-108 Ichlhyosauria Platypterygidae Platypterygius hercyiiicus b/d/e/h/i 6 17 EU-480 S-e-f MR-882 Ichthyosauria indet. indet. indet. e 5 82; 176 EU-480 S-e-f MR-884 Ichlhyosauria indet. indet. indet. e 5 82; 176 EU-480 S-e-f MR-875 Ichthyosauria indet. indet. indet. e 5 82; 176 TABLE G.2 Marine Reptile Database Taxa

Loc Map MMR Order Family Genus Spccics Complete Reference ID Interval ID * * \faltr\n

EU-480 s-c-r MR-773 Plesiosauria Cr>'ptoclididae C'ulyinbosaurus trochantcrius c 0 8I;40 EU-480 S-c-f MR-780 PIcsiosauria Cryptoclididae Cryptocleidus afT. richardsoni h 0 81; 40 EU-480 S-c-f MR-772 Plesiosauria indct. indet. indet. e/h 0 81; 40 ElJ-480 S-e-f MR-80() Plesiosauria Pliosauridae Liopleurodon macromerus e 0 81; 291 EU-480 $-e-r MR-792 Plesiosauria Pliosauridae Pliosaurus brachydcirus d/e 0 81 EU-481 c-e-f MR-774 Plesiosauria Ciyptoclididae Colymbosaurus trochantcrius S 0 8I;40 EU-482 c-e-1' MR-775 PIcsiosauria Cryptoclididae Colymbosaurus trochantcrius S 0 8I;40 EU-483 e-e-f MR-776 PIcsiosauria Cryptoclididae Colymbosaurus trochantcrius s 0 8I;40 EU-484 $-d-c MR-782 PIcsiosauria indet. indei. indet. i 0 81 EU-485 S-d-e MR-783 PIcsiosauria indet. indet. indct. d 0 81 EU-486 $-e-r MR-785 Plesiosauria indet. indet. indct. d 0 81 EU-486 $-e-f MR-803 Plesiosauria Pliosauridae Liopleurodon ?macrometus d 0 81; 291 EU-486 S-c-f MR-793 Plesiosauria Pliosauridae Pliosaurus brachydcirus b 0 81 EU-486 $-c-f MR-807 Plesiosauria Pliosauridae Pliosaurus SP S 0 81 EU-487 S-c-f MR-891 Ichlhyosauria Ichthyosauridae Ophthalmosaurus sp. e/z 6 82 EU-487 S-e-f MR-787 PIcsiosauria indet. indet. indet. f 0 81 EU-488 e-c-f MR-789 Plesiosauria indet. indet. indet. e 0 81 EU-489 s-e-r MR-801 Plesiosauria Pliosauridae Liopleurodon macromerus e 0 81; 291 EU-490 S-e-f MR-876 Ichthyosauria indet. indet. indet. e 5 82; 176 EU-490 S-c-f MR-810 PIcsiosauria Pliosauridae Pliosaurus sp. S 0 81 EU-491 S-e-f MR-8IS Ichthyosauria indet. indct. indet. e/f/7. 5/6 218 EU-507 S-c-d MR-834 Ichthyosauria Ueptopterygidac Excalibosaurus costini b/c/d/c/f/h/i 6 178 EU-512 S-d-c MR-877 Ichthyosauria indet. indet. indet. e 6 82; 176 EU-513 S-e-f MR-886 Ichthyosauria indet. indct. indet. c 5 82; 176 EU.514 S-e-f MR-870 Ichthyosauria indet. indet. indct. e 5 82; 176 EU-515 S-h-g MR-894 Ichthyosauria Platypterygidae Platyptcrygius "campylodon" d 2 82; 173 EU-516 S-e-f MR-890 Ichthyosauria Ichthyosauridae Ophthalmosaurus sp. h 5 82 EU-516 S-e-f MR-909 Ichthyosauria indet. indet. indet. z 5 82 EU-527 S-f-g MR-910 Plesiosauria Pliosauridae indet. indet. d 2 309 EU-528 S-i-h MR-911 Plesiosauria Pliosauridae indet. indet. d 2 309 EU-53 h-h-h MR-II3 Ichthyosauria Platyptet>'gidae Platyptcrygius f P. kiprijanoffi b/c/d/c/f 6 17 EU-530 S-i-h MR-913 Plesiosauria Pliosauridae indet. indct. c 0 309 EU-54 S-c-d MR-114 Ichthyosauria Ichthyosauridae Ichthyosaurus indct. b/c/c/f/h/I 6 91 EU-541 S-c-d MR-959 Ichthyosauria indct. indet. indct. d/e 6 134 EU-542 S-a-b MR-960 Ichthyosauria Shastasauridae Cymbospondylus sp. b/cyd/e/f/h/i 6 142;48 EU-543 d-d-c MR-961 Plesiosauria indet. indet. indct. d/e/f 6 263; 264 EU-55 S-c-d MR-496 Ichthyosauria I.eptopterygidBC Eurhinosaurus longirostris b 6 177; 184 EU-55 S-c-d MR-II5 Ichthyosauria Stenoptci>'gidae Stenopterygius mcgaccphalus b/c/d/e/f/g/h 7 129 TABLE G.2 Marine Reptile Database Taxa

Loc Map MMR Order Tamily Genus Snccies Complete .. Reference ID Interval ID ' ' vation

/i l:U-55 $-c-d MR-II7 Ichthyosauria Stenoplerygidae Slenoplerygius quadriscissus b/c/e/f/li/i 7 129 EU-553 $-i-h MR-I0I3 Squamata Mosasauridae Leiodon anceps S 0 27 EU-553 $-i-h MR-I0I4 Squamaia Mosasauridae Mosasaums sp. S 0 27 EU-554 $-l-h MR-1003 Squamata Mosasauridae Plalecarpus sp. s 0 27 EU-555 $-l-h MR-1004 Squamata Mosasauridae llainosaurus? sp. e 5 18; 27 EU-556 $-i-h MR-1045 Plesiosauria Pliosauridae indel. indct. S 0 27 EU-557 $-i-h MR-IOOl Squamata Mosasauridae llainosaurus bemardi b/c/d 5 18; 27 EU-557 $-i-h MR-1002 Squamata Mosasauridae Plalecarpus sonienensis S 0 27 EU-558 $-i-h MR-1046 Plesiosauria Pliosauridae indet. indel. e 0 22; 27 EU-558 $-i-h MR-IOOO Squamata Mosasauridae llainosaurus sp. S 0 27 EU-558 $-i-h MR-999 Squamata Mosasauridae Mosasaurus lonzeensis S 0 27 EU-559 $-i-h MR-1007 Squamata Mosasauridae Prognathodon giganteus s 0 27 EU-56 $-c-d MR-491 Ichthyosauria Leptopterygidae Eurhinosaurus longirostris a 7 177; 184 EU-56 $-c-d MR-490 Ichthyosauria Slenoptetygidae Stenopterygius macrophasma a 7 177 EU-56 $-c-d MR-489 Ichthyosauria Stenoplerygidae Slenoplerygius megacephalus a 7 177 EU-56 $-c-d MR-II6 Ichlhyosauria Stenoplerygidae Slenoplerygius quadriscissus b/c/e/l"/i 7 129 EU-56 $-c-d MR-122 Ichlhyosauria Stenoplerygidae Slenoplerygius quadriscissus b/c/e/f/h/i 7 129 EU-56 $-c-d MR-119 Ichlhyosauria Stenoplerygidae Slenoplerygius quadriscissus b/c/e/f/h/i 7 129 EU-56 $-c-d MR-488 Ichlhyosauria Stenoplerygidae Slenoplerygius quadriscissus a 7 177 EU-56 $-c-d MR-120 Ichthyosauria Stenopteiygidae Slenoplerygius quadriscissus b/c/e/f/hyi 7 129 EU-56 $-c-d MR-492 Ichlhyosauria Temnodontosaurid Temnodontosaurus burgundiae b 6 177; 186 dc EU-560 $-l-h MR-1005 Squamata Mosasauridae Leiodon compressidcns S 0 27 EU-561 $-i-h MR-IOlO Squamata Mosasauridae Plalecarpus sp. S 0 27 EU-562 S-l-h MR-ID2I Squamata Mosasauridae Mosasaurus sp. s 0 27 EU-563 S-i-h MR-I006 Squamata Mosasauridae Leiodon compressidcns 0 27 EU-564 $-i-h MR-1042 Plesiosauria Elasmosauridae indel. indel. s 0 27 EU-565 $-i-h MR-I0I9 Squamata Mosasauridae Leiodon sp. s 0 27 EU-565 S-i-h MR-I0I8 Squamata Mosasauridae Mosasaurus sp. 0 27 EU-565 $-i-h MR-1020 Squamata Mosasauridae Plalecarpus sp. s 0 27 EU-566 S-i-h MR-1008 Squamata Mosasauridae Leiodon anceps s 0 27 EU-566 S-i-h MR-1009 Squamata Mosasauridae Mosasaurus sp s 0 27 EU-567 S-i-h MR-1023 Squamata Mosasauridae Mosasaurus sp. s 0 27 EU-S68 $-i-h MR-IOII Squamaia Mosasauridae Globidens dakoiaensis s 0 27 EU-568 S-i-h MR-I0[2 Squamata Mosasauridae Leiodon sp. s 0 27 EU-569 S-i-h MR-I022 Squamata Mosasauridae Mosasaurus sp. s 0 27 EU-57 S-c-d MR-495 Ichlhyosauria l.eploplerygtdae l-.urhinosaurus longirostris a 7 177; 184 u> u> TABLE G.2 Marine Reptile Database Taxa

1.0C Map MMR ^ . Family Genus Species Complete . ' Reference ID Interval ID ' ' vation

IUJ-57 $-c-d MR-494 Ichlhyo&auria Stenopterygidae Stenopterygius cuneiceps a 7 177 EU-57 $-c-d MR-493 IchthyosBuiia Stenoptciygidae Stenopleiygius hauflianus aA> 6/7 177 EU-57 $-c-d MR-118 Ichthyosauria Stenopterygidae Stenopterygius quadriscissus b/c/c/r/h/i 7 129 l:U-570 j-j-h MR-1024 Squamala Mosasauridae Carinodens belgicus S 0 27 ElI-570 j-j-h MR-1025 Squaniata Mosasauridae Carinodens Traasi s 0 27 EU-570 j-j-h MR-1026 Squaniata Mosasauridae llainosaunis bcmardi a 7 147; 27 EU-570 j-j-h MR-llll Squamala Mosasauridae llalisaurus ortliebi b 5 151 F.U-570 j-j-h MR-I03I Squamala Mosasauridae llalisaurus sp. S 0 27 EU-570 j-j-h MR-1027 Squamala Mosasauridae Mosasaurus lemnionieri s 0 27 EU-570 j-j-h MR-1028 Squaniata Mosasauridae I'lioplatecarpus houzeaui $ 0 27 EU-570 j-j-h MR-1029 Squaniata Mosasauridae Prognathodon giganteus $ 0 27 EU-570 j-j-h MR-1030 Squamata Mosasauridae Prognathodon solvayi $ 0 27 EU-571 j-j-h MR-1043 Plesiosauria Elasmosauridae indet. indet. e 0 27 EU-572 j-j-h MR-1044 Plesiosauria Elasmosauridae indet. indet. d 0 200;27 EU-572 j-j-h MR-1032 Squamata Mosasauridae Carinodens Traasi S 0 27 EU-572 j-j-h MR-1035 Squaniata Mosasauridae Leiodon sectorius $ 0 27 EU-572 j-j-h MR-1033 Squamata Mosasauridae Mosasaurus hofTmanni q 5 150;27 EU-572 j-j-h MR-1034 Squamala Mosasauridae Plioplatecaqms maibhi s 0 27 EU-573 j-j-h MR-1037 Squamata Mosasauridae Leiodon mosasauroides s 0 27 EU-574 j-j-h MR-1038 Squamata Mosasauridae Leiodon anceps s 0 27 EU-574 j-j-h MR-1039 Squamata Mosasauridae Mosasaurus sp. s 0 27 EU-575 j-j-h MR-1040 Squamata Mosasauridae indet. indet. s 0 27 EU-576 j-j-h MR-1036 Squamata Mosasauridae Carinixlens fraasi s 0 27 EU-577 $-l-h MR-I04I Plesiosauria Elasmosauridae indet. indet. e 5 22; 27 EU-577 $-i-h MR-1069 Plesiosauria indet. indet. indet. e/h/i 5 22 EU-S8 $-c-d MR-I2I Ichthyosauria Stenopterygidae Stenopterygius quadriscissus b/c/e/r/h/i 7 129 EU-582 $-d-c MR-1055 Ichthyosauria indet. indet. indet. e/r 5 7 EU-S84 h-h-h MR-1058 Ichthyosauria Platypterygidae Platypterygius "campylodon" e 4 19 EU-59 $-c-d MR-123 Ichthyosauria Stenopterygidae Stenopterygius megacephalus b/c/e/r/i 6 177, 129 EU-60 S-h-g MR-124 Ichthyosauria Platypterygidae Platypterygius sp. b/c 2 23 EU-609 S-b-c MR-1108 Plesiosauria Pistosauridae Pistosaurus longaevus h 5 74 EU-61 $-c-d MR-125 Ichthyosauria 1'emnodontosaurid Tem nodontosaurus platyodon b/c/d/e/f/h/i 6 175;115 ae EU-61 $-c-d MR-308 Plesiosauria Pliosauiidae indet. indet. e 4 117 EU-610 j-j-h MR-1109 Squamata Mosasauridae Mosasaurus hoflmanni b/c/d 5 276 EU-611 S-a-a MR-1112 Ichthyosauria incertae sedis Svalbardosaurus crassidens d 5 166 EU-611 S-a-a MR-1113 Ichthyosauria indet. indet. indet. b 5 166 EU-612 $-a-c MR-1114 Plesiosauria indet. indet. indet. e/f 5 72 u> u> 00 TABLE G.2 Marine Reptile Database Taxa

Loc Map MMR Order l-'aniily Genus Spccics Complcic .. " Rererencc ID 1Interval ID * * vntmn

1:IJ-6I3 $-b-c MR-IIIS Ichthyosauria indet. indet. indei. e/r/i 6 72 EU-614 $-a-b MR-III6 Ichlhyosauria indet. indet. indet. e/f 5 72 l-lJ-615 $-a-b MR-III7 Ichthyosauria indei. indet. indei. e/r 5 72 l-U-616 $-a-b MR-III8 Ichthyosauria incertae sedis Pessosaurus sp. e/r 5 72; 48 EU-617 S-a-b MR-III9 Ichthyosauria indet. indet. indet. e/r 5 72 EU-618 $-a-b MR-1120 Ichthyosauria indet. indet. indei. e/r 5 72 l-U-619 $-a-b MR-II2I Ichthyosauria indet. indet. indet. e/r 5 72 l-U-620 $-a-b MR-1122 Ichlhyosauria indet. indet. indet. e/r 5 72 EU-621 $-a-b MR-II23 Ichlhyosauria indet. indet. indet. e/r 5 72 EU-622 $-a-b MR-1124 Ichlhyosauria Mixosauridae Mixosaurus sp. d/e/r 5 72 EU-623 S-a-a MR-1209 Ichlhyosauria indet. Orippia longirostris b/c/d/e/r/h/i 6 167 EU-623 $-a-a MR-1125 Ichthyosauria indet. indet. indet. e/r 5 72 EU-624 S-a-a MR-1126 Ichthyosauria indet. indet. indet. e/r 5 72 EU-625 S-a-a MR-1208 Ichthyosauria indei. Grippia longirostris b/c/d/e/r/h/i 6 167 EU-625 S-a-a MR-1127 Ichthyosauria indet. indet. indet. e/r 5 72 EU-626 $-a-a MR-1128 Ichthyosauria indet. indet. indet. e/r 5 72 EU-627 $-a-a MR-1129 Ichthyosauria indet. indet. indet. e/r 5 72 EU-631 $-c-d MR-1135 Ichthyosauria Leptopleo'gidae Eurhinosaurus longirostris hide 6 143 EU-632 $-d-e MR-II36 Plesiosauria Pliosauridae Simolesles keileni c/d/h 6 116 EU-633 S-l-h MR-1137 Squaniata Mosasauridae Prognathosaurus sp. c/d 5 326 EU-634 j-j-h MR-1138 Squamata Mosasauridae llainosaurus bemardi b/c/d/e 6 147 EU-637 $-b-c MR-1152 Ichthyosauria indet. indei. indet. e/h 5 21 ElJ-637 $-b-c MR-1153 Plesiosauria indet. indet. indet. d 5 21 EU-639 $-b-c MR-1155 Ichthyosauria indei. indet. indet. d 3 21 EU-639 $-b-c MR-1156 Plesiosauria indet. indet. indet. d 3 21 EU-640 $-b-c MR-n57 Ichthyosauria I.eptoptcrygiidae indet. indet. e 3 21 ElJ-640 $-b-c MR-1158 Plesiosauria indei. indet. indet. d/e 3 21 EU-641 $-a-b MR-1164 Plesiosauria Pistosauridae indet. indet. b/c/d 6 261; 3 UU-654 $-b-c MR-1188 Ichthyosauria indei. indet. indet. e/r 5 321 EU-654 $-b-c MR-1189 Plesiosauria indet. indet. indet. r/g/h/i 5/6 321 r:l)-66 $-a-b MR-U78 Ichlhyosauria Mixosauridae Mixosaurus comalianus q 7 141 Ell-66 $-a-b MR-133 Ichlhyosauria Shasiasauridae Cymbospondylus buchseri b/c/d/e/r/g/h 6 255 /i EU-663 $-a-b MR-1369 Ichlhyosauria indet. indet. indet. s 0 48 EU-665 $-e-f MR-1205 Plesiosauria Pliosauridae Pliosaurus brachyspondylus c 5 25 EU-666 $-c-d MR-1206 Ichthyosauria Stenopterygidae cr. Slenopterygius sp b/d 6 258 EU-667 d-d-e MR-1207 Plesiosauria Elasmosauridae Muraenosaunis leedsii e/h 5/6 26 EU-668 $-a-b MR-I2I2 Ichthyosauria incertae sedis Pessosaurus pularis c/d/h 5 169

SO TABLE G.2 Marine Reptile Database Taxa

Loc Map MMR Order l-amily Genus Specics Complete . ' Reference ID Interval ID l:U-668 $-a-b MR-I2II Ichlhyosauria incerlae sedis Phalarodon cf P. fraasi c/d 6 169 (:U-668 $-a-b MR-1213 Ichthyosauria Mixosauridae Mixosaurus nordenskioeldii e/f/h 5/6 169 l:U-669 $-c-d MR-I2I4 Pleslosauria Elasniosauridae cl'. Microcleidus sp. h 5 318 EU-674 d-d-e MR-I23I Plesiosauria indet. indet. indet. e/g/i 6 69 EU-679 $-i-h MR-1247 Squaniata Mosasauridae Leiodon cf L. anceps d 4 225 KU-679 $-i-h MR-1246 Squamala Mosasauridae Plioplatecarpus? sp. d 4 225 l-U-681 $-c-d MR-1252 Plesiosauria Plesiosauridae Plesiosaurus toiimcmirensis a 6 267 ElJ-689 r-f-B MR-1260 Ichthyosauria indet. indet. indet. e 4 294 EU-690 f-f-g MR-I26I Ichthyosauria indet. indet. indet. e 4 294 EU-703 e-e-f MR-1294 Ichthyosauria Ichthyosauridae Ichthyosaurus sp. S 6 30 EU-703 e-«-f MR-1295 Ichthyosauria indet. indet. indet. S 6 176; 30 EU-703 e-e-f MR-1296 Plesiosauria Pliosauridae Liopleurodon sp. d 4 30 EU-704 j-j-h MR-1297 Squamata Mosasauridae Mosasaurus sp. b/d 5 289 EU-705 j-j-h MR-1298 Squaniata Mosasauridae Mosasaurus sp. d 4 289 EU-713 $-c-d MR-I3I4 Plesiosauria indet. indet. indet. i 5 133 EU-72 $-e-f MR-139 Ichthyosauria Ichthyosauridae Brachypterygius extremus b/c/d/df/h 6 176;187 EU-720 S-e-f MR-1327 Ichthyosauria Ichthyosauridae Ophihalmosaurus sp. e/h/z 5 302 EU-733 S-S-d MR-1347 Ichthyosauria Ichthyosauridae Ichthyosaurus sp. d 4 199 EU-735 $-b-c MR-1349 Ichthyosauria indet. ?Shonisaurus sp. d/e/r 0 48 EU-742 S-a-c MR-1358 Ichthyosauria Shastasauridae Shastasaurus sp. $ 0 48; 182 EU-743 $-$-b MR-1359 Ichthyosauria Shastasauridae ?Shastasaurus sp. S 0 48 EU-744 $-a-b MR-1362 Ichthyosauria indet. indet. indet. e 4 48 EU-745 $-a-b MR-1364 Ichthyosauria incertae sedis Pessosaurus sp. e 0 48 EU-745 $-a-b MR-1365 Ichthyosauria Shastasauridae Shastasaurus sp. e 0 48 EU-746 $-a-b MR-1366 Ichthyosauria Shastasauridae Shastasaurus sp. e 0 48 EU-749 S-a-b MR-1370 Ichthyosauria indet. indet. indet. e 0 48 EU-750 $-a-b MR-1371 Ichthyosauria Mixosauridae Mixosaurus comalianus a 7 232;48 EU-751 $-a-b MR-1383 Ichthyosauria Mixosauridae indet. indet. S 0 48 EU-751 $-a-b MR-1372 Ichthyosauria Shastasauridae Cymbospondylus sp e/f 0 48 EU-752 $-a-b MR-1386 Ichthyosauria incertae sedis Pessosaurus sp. e 0 48 EU-752 $-a-b MR-1377 Ichthyosauria Mixosauridae Mixosaurus atavus $ 0 48 EU-752 $-a-b MR-1376 Ichthyosauria Mixosauridae Mixosaurus sp h 0 48 EU-752 $-a-b MR-1389 Ichthyosauria Shastasauridae '.'Shastasaurus sp. e 0 48 EU-752 $-a-b MR-1375 Ichthyosauria Shastasauridae Cymbospondylus sp. e/f 0 48 EU-752 $-a-b MR-1388 Ichthyosauria Shastasauridae indet. indet. e 0 48 EU-759 S-a-b MR-1395 Ichthyosauria Shastasauridae Cymbospondylus sp. e/f 6 256 EU-76 h-h-h MR-153 Ichthyosauria Platypteo'gidac Platypterygius "campylodon" c/d 2 173; 243;19 EU-79 $-h-g MR-158 Ichthyosauria indet. indet. indet. b/c/d 3 79 u> O TABLE G.2 Marine Reptile Database Taxa

Loc Map MMR Order Family Cienus Species Complele feser- Refjfenge ID Interval ID ' ' vation

EU-79 $-h-g MR-157 Ichlhyosauria Plalyplerygidae Plalyplerygius sp. b/c 3 79 EU-80 $-e-f MR-159 Ichlhyosauria indei. indel indel. e/f/g/lj/i 5/6 78 l-U-81 c-c-d MR-160 Ichlhyosauria Ichlhyosauridae Ichthyosaurus communis e 5 76; 174 EU-81 c-c-d MR-664 Ichlhyosauria Slenoplerygidac '.'Stenopieiygius megalorhinus b/d 5 76; 177 EU-8II $-f-g MR-1459 Plesiosauria Elasmosauridae indel. indet. e/i 3 100 ElJ-812 c-c-d MR-1460 Ichlhyosauria Leploplerygidae I.cptonectes leniurosiris h/i 5 114;186 EU-814 S-a-b MR-1462 Ichlhyosauria indel. indet. indel. c 6 38 EU-815 $-c-f MR-1463 Ichlhyosauria indel. indel. indel. b/c/d/c 6 92 EU-85 s-e-r MR-164 Plesiosauria Pliosauridae Peloneusles sp. c/d 4 163 EU-86 $-e-f MR-I6S Plesiosauria Pliosauridae Pliosaurus cf. P. andreusi b 4 163 EU-87 $-i-h MR-166 Plesiosauria Pliosauridae indel. indel. h 3 228; 309 EU-88 $-l-h MR-167 Plesiosauria Elasmosauridae indel. indel. z 3 228; 309 EU-88 $-i-h MR-t68 Squamala Mosasauridae Leiodon sp. d 2 228 EU-89 $-i-h MR-169 Plesiosauria Plesiosauridae indel. indet. h 3 228 EU-89 $-i-h MR-175 Plesiosauria Pliosauridae indet. indet. e 4 309;230 EU-89 $-i-h MR-357 Plesiosauria Pliosauridae indet. indel. d/c/h/i 0 309; 229 EU-89 $-i-h MR-I0I6 Squamala Mosasauridae Leiodon sp. S 0 27 EU-89 $-i-h MR-1243 Squamala Mosasauridae Mosasaurus cf. M. holTmanni d 4 225 EU-89 $-i-h MR-I0I3 Squamala Mosasauridae Mosasaurus sp. S 0 27 EU-89 $-i-h MR-1244 Squamala Mosasauridae Mosasaurus sp. d 4 225 EU-89 $-i-h MR-1245 Squamala Mosasauridae Plalecarpus cf P. somenensis d 4 225 EU-89 $-i-h MR-1017 Squamala Mosasauridae Platecarpus sp. $ 0 27 EU-90 $-$-h MR-176 Plesiosauria Pliosauridae indet. indet. e 4 309; 230 EU-90 S-S-h MR-170 Plesiosauria Pliosauridae indel. indel. e 3 228; 309 EU-95 S-i-h MR-177 Plesiosauria indel. indet. indet. e/i/z 2 230 EU-96 S-d-e MR-178 Plesiosauria indet. indel. indel. e/i 6 113 NA-I $-a-c MR-I Ichlhyosauria Shonisaurldae Shonisaurus popularis q/z 6 52; 127 NA-IO S-i-h MR-13 Plesiosauria Polycolylidae Dolichorhynchops obomi z 0 233;56 NA-IO $-i-h MR-12 Squamala Mosasauridae Platecarpus ictericus z 0 233 NA-II j-j-h MR-1470 Plesiosauria Elasmosauridae indel. indel. S 0 251 NA-II j-j-h MR-14 Plesiosauria indel. indel. indet. z 0 233;251 NA-ll j-j-h MR-1472 Squamala Mosasauridae cf. Mosasaurus sp. $ 0 251 NA-II j-j-h MR-I47I Squamala Mosasauridae Plioplatecarpus sp S 0 251 NA-13 S-a-c MR-16 Ichlhyosauria indel. indel. indel. q/z 1/2/3/ 188 4/5/6/ 7 NA-I 4 b-b-c MR-17 Ichlhyosauria Shaslasauridae Shaslasaunis neoscapularis b/c/d/e/f/g/h 7 188 /i TABLE G.2 Marine Reptile Database Taxa

I.OC Map MMR Order Family Genus Spccics Complete .. " Reference ID Interval ID ' ' vnimn

NA-15 b-b-c MR-18 Ichthyosauria indet. lludsonelpidia brevirostris q 6/7 188 NA-16 $-$-c MR-19 Ichlhyosauria indet. indet. indet. b/i/c/f/h/z/d/ 6/7 188 c NA-17 b-b-c MR-20 Ichthyosauria indet. indet. indet. i/e/r 6 188 NA-18 b-b-c MR-21 Ichthyosauria Ichthyosauridae Ichthyosaurus janiceps b/h/l 6 185;188 NA-243 $-e-r MR-410 Plesiosauria indet. indet. indet. e/h/i 0 229 NA-249 j-j-h MR-416 Plesiosauria Hlasmosauridae Lcurospondylus ultimus e/h/i 0 229 NA-250 S-i-h MRm7 Plesiosauria Elasmosauridae Alzadasauius pembenoni a 0 307;229 NA-251 $-i-h MR-418 Plesiosauria Elasmosauridae Hlasmosaurus platyurus a 0 307; 229 NA-252 $-l-h MR-419 Plesiosauria Hlasmosauridae llydralmosaurus serpcntinus e/h/i 0 307; 229 NA-253 S-i-h MR-420 Plesiosauria i-lasmosauridae Alzadasauius kansasensis e/h/i 0 229 NA-254 S-i-h MR-421 Plesiosauria Hlasmosauridae Styxosaurus browni a 0 229 NA-255 S-i-h MR-424 Plesiosauria I-lasmosauridae '.'Hlasmosaurus sp. e/h/i 0 309; 229 NA-255 S-i-h MR-423 Plesiosauria Hlasmosauridae indet. indet. e/h/i 0 309; 229 NA-255 S-i-h MR-422 Plesiosauria Hlasmosauridae indet. indet. b/c/e 0 309; 229 NA-255 S-i-h MR-431 Plesiosauria Polycotylidae Dolichorhynchops osbomi a/q 0 229; 56 NA-256 i-i-h MR-425 Plesiosauria Hlasmosauridae indet. indet. e/h/i 0 307; 309; 229 NA-257 S-i-h MR-426 Plesiosauria Hlasmosauridae Libonectes morgani a 0 229; 57 NA-258 S-h-h MR-427 Plesiosauria Hlasmosauridae Thalassomedon haningtoni a 7 307; 229 NA-259 S-h-h MR-428 Plesiosauria Hlasmosauridae Alzadasauius riggsi e/h/i 0 229 NA-260 S-S-h MR-429 Plesiosauria Pliusauroidea indet. indet. e 0 309; 229 NA-261 S-i-h MR-430 Plesiosauria Polycotylidae Polycotylus latipinnus aJdd'x 0 229; 56 NA-262 S-i-h MR-432 Plesiosauria Polycotylidae Trinacromerum bentonianum c/e/i 0 229; 56 NA-263 S-i-h MR-433 Plesiosauria Polycotylidac 1'rinacromeium bentonianum b/e/h 0 229; 56 NA-264 S-h-h MR-434 Plesiosauria Pliosauridae Brachauchenius lucasi b/c/h 0 229 NA-265 S-i-h MR-435 Plesiosauria Polycotylidae Trinacromerum kirki e/f/g/h/i 6 229; 56 NA-266 S-i-h MR-437 Plesiosauria Polycotylidae indet. indet. e 0 229 NA-265 S-i-h MR-436 Plesiosauria Polycotylidae indet. indet. e 0 229 NA-267 S-S-h MK-438 Plesiosauria Polycotylidae indet. indet. d 0 229 NA-268 i-i-h MR-439 Plesiosauria Polycotylidae indet. indet. e 0 229 NA-269 j-j-h MR-440 Plesiosauria indet. indet. indet. e/i 0 307; 229 NA-270 S-i-h MR-441 Plesiosauria indet. indet. indet. e 0 229 NA-271 S-h-g MR-442 Plesiosauria Hlasmosauridae indet. indet. e 0 229 NA-272 S-g-6 MR-443 Plesiosauria Hlasmosauridae indet. indet. e 0 229 NA-272 S-B-g MR-444 Plesiosauria Polycotylidac indet. indet. e 0 229 NA-273 S-d-e MR-I53I Ichlhyosauria Ichthyosauridae Baptanodon sp. b/c/d/e/f/i 0 112 NA-273 S-d-e MR-445 Plesiosauria indet. indet. indet. c/d/e/i 0 229 NA-274 S-d-e MR-1530 Ichlhyosauria Ichthyosauridae Baptanixlun sp. b/c/d/e/h/i 0 112 TABLE G.2 Marine Reptile Database Taxa

Loc Map MMR Order Family Genus Species Complete , . Reference ID Interval ID

NA-274 $-d-c MR-446 Plesiosauria indet. indet. indet. e/i 0 229 NA-275 $-d-c MR-447 PIcsiosauria Pliosauridae Megalneusaurus rex e/h/i 0 229 NA-276 j-j-h MR-448 Plesiosauria Elasmosauridae indet. indet. e 0 229 NA-277 $-$-g MR-449 Plesiosauria Elasmosauridae indet. indet. d 0 229 NA-278 $-i-h MR-450 Plesiosauria Hlasmosauridae Aphrosaurus lurlongi e/h/i 0 307; 229 NA-278 S-i-h MR-451 Plesiosauria Hlasmosauridae Fresnosaunis drescheri h/i 0 229 NA-278 $-i-h MR-452 Plesiosauria l-lasmosauridae Ilydrotherosaurus alexandrae a 0 307;229 NA-278 S-i-h MR-453 Plesiosauria Hlasmosauridae Morenosaurus stocki e/h/i 0 307; 229 NA-279 e-c-f MR-454 Plesiosauria indet. indet. indet. e 0 229 NA-286 $-$-h MR-474 Plesiosauria Hlasmosauridae indet. indet. e/i 0 247 NA-286 $-$-h MR-475 Plesiosauria Hlasmosauridae indet. indet. e/i 3 247 NA-286 $-$-h MR-472 Plesiosauria Polycotylidae Dolichorhynchops osbomi e/h 6 247;57 NA-286 $-$-h MR-473 Plesiosauria Polycotylidae indet. indet. e 3 247 NA-286 $-$-h MR-471 Squamaia Mosasauridac Platecarpus ictericus bide 3 247 NA-287 j-j-h MR-476 Plesiosauria indet. indet. indet. e/i 6 247 NA-288 $-i-h MR-1599 Plesiosauria Hlasmosauridae '.'HIasmosaurus sp. e/h/i 6 207 NA-288 $-i-h MR-1598 Plesiosauria Hlasmosauridae cf. Alzadasaurus pembertoni e/h/i 5/6 207 NA-288 $-i-h MR-1600 Plesiosauria Hlasmosauridae indet. indet. e/h 6 207 NA-288 $-i-h MR-1597 Plesiosauria Polycotylidae indet. indet. biddldi 4 207 NA-288 $-l-h MR-1594 Plesiosauria Polycotylidae Trinacromerum bonneri a/q 6 207; 2 NA-288 $-i-h MR-1595 Plesiosauria Polycotylidae 1 rinacromerum cf. T. kirki a 6/7 207 NA-288 S-i-h MR-1596 Plesiosauria Polycotylidae 1'rinacromerum sp. b/c/d/e/h/i 4 207 NA-288 S-i-h MR-1590 Squamata Mosasauridae Clidastes propyihon b/c/d/e 6 207 NA-288 S-i-h MR-477 Squamata Mosasauridae Hainosaurus pembinensis b/c/d/e/h/i 6 206 NA-288 S-i-h MR-1589 Squamata Mosasauridae indet. indet. e 4/5 207 NA-288 S-i-h MR-1592 Squamata Mosasauridae indet. indet. li/i 4/5 207 NA-288 S-i-h MR-1593 Squamata Mosasauridae indet. indet. e/f/i/z 4 207 NA-288 S-i-h MR-1587 Squamata Mosasauridae Platecarpus somenensis q 4/5/6 207 NA-288 S-i-h MR-1588 Squamata Mosasauridae Platecarpus sp. b/h/i 4/5 207 NA-288 S-i-h MR-1586 Squamata Mosasauridae Platecarpus tympaniticus q 4/5/6 207 NA-288 S-i-h MR-1591 Squamaia Mosasauridae Tylosaurus prorigor c/e/h 5 207 NA-289 S-a-a MR.478 Ichthyosauria inceitae sedis IJtatsusaurus sp. b/c/d 6 209 NA-289 S-a-a MR-10% Ichthyosauria indet. Grippia cf. U. longirosiris e/f/g/h/i/z 6 47; 39 NA-290 S-i-h MR-479 Plesiosauria Hlasmosauridae indet. indet. S 0 208 NA-297 j-j-h MR-486 Plesiosauria Hlasmosauridae '?Leurospondylus sp e 3 253 NA-318 j-j-h MR-537 Squamata Mosasauridae llalisaurus platyspondylus b 4 13 NA-318 j-j-h MR-538 Squamata Mosasauridae Mosasaurus conondon e 3 13 NA-319 S-i-h MR-540 Squamata Mosasauridae Platecarpus sp. ddldf 5 157 U) U) TABLE G.2 Marine Reptile Database Taxa

Loc Map MMR Preset- Order Tamily Genus Specics Complclc . Kererence ID Interval ID ' ' vaiion

NA-321 $-i-h MR-542 Squamata Mosasauridac indet. indct. c 3 162 NA-322 $-l-h MR-543 Squaniaia Mosasauridae indct. indet. c 4 162 NA-323 S-l-h MR-544 Squamata Mosasauridae indct. indcl. e/z 4 162 NA-324 j-j-h MR-545 Plesiosauria ?Pliosauroidea indct. indcl. d/c/f 4 309; NA-325 j-j-h MR-546 I'leslosauria indet. indet. indct. die 3 223 NA-327 j-j-h MR-S48 Pleslosauria indct. indct. indct. i 3 314 NA-327 j-j-h MR-1236 Squamata Mosasauridac Mosasaurus maximus $ 0 71 NA-327 j-j-h MR-1237 Squamata Mosasauridae I'lioplatecarpus dcprcssus S 0 71 NA-327 j-j-h MR-1238 Squamata Mosasauridae Prognathodon sp. S 0 71 NA-328 $-l-h MR-549 Squamata Mosasauridae Clidastes iguanavus c 0 248 NA-329 S-l-h MR-550 Squamata Mosasauridae Clidastes stembcrgi a 7 248 NA-33 $-a-c MR-57 Ichthyosauria indet. indet. indet. e 0 290 NA-33 $-a-c MR-58 Ichthyosauria Mixosauridae Mixosaurus sp. d 1 290 NA-330 $-i-h MR-551 Squamata Mosasauridae Clidastes liodonlus b/c/d/z 0 248 NA-330 $-i-h MR-578 Squamata Mosasauridae llalisaurus onchognathus b/e 0 248 NA-330 $-i-h MR-1475 Squamata Mosasauridae Platecarpus lympaniticus S 0 M.J. NA-331 $-i-h MR-552 Squamata Mosasauridae Clidastes propyihon a 7 248 NA-332 j-j-h MR-553 Squamata Mosasauridae Mosasaurus conodon b/c/d/e/h/i 0 248 NA-333 j-j-h MR-554 Squamata Mosasauridae Mosasaurus missouriensis b/e/h 0 248 NA-334 j-j-h MR-555 Squamata Mosasauridae Mosasaurus dekayi d 1 248 NA-335 j-j-h MR-556 Squamata Mosasauridae Mosasaurus maximus b/c/d/e 0 248 NA-336 $-i-h MR-557 Squamata Mosasauridae Amphekepubis johnsoni e/h/i 0 248 NA-337 j-j-h MR-558 Squamata Mosasauridae Leiodon sectorius b/c/d/e 0 248 NA-338 $-l-h MR-1076 Squamata Mosasauridae Clidastes propyihon q 5 249 NA-338 $-i-h MR-559 Squamata Mosasauridae Globidcns alabamaensis b/c/d/c/z 0 248; NA-338 S-i-h MR-1075 Squamata Mosasauridae indct. indcl. b/e 5 249; NA-338 $-i-h MR-1084 Squamata Mosasauridae indct. indcl. i 5 249 NA-338 $-i-h MR-1087 Squamata Mosasauridac indcl. indet. c 5 249 NA-338 $-i-h MR-1080 Squamata Mosasauridae Platecarpus sp. b/c/d/c/f/i 6 249 NA-338 $-i-h MR-I08I Squamata Mosasauridae Prognathodon sp. h/i 5 249 NA-338 $-i-h MR-1085 Squamata Mosasauridae Tylosaurus sp. b/c/i 6 249 NA-339 $-i-h MR-560 Squamata Mosasauridae Globidcns alabamaensis b 0 248 NA-34 $-a-c MR-59 Ichthyosauria indet. indct. indcl. e 6 290 NA-340 S-i-h MR-561 Squamata Mosasauridae Globidens alabamaensis c/d 0 248 NA-341 $-i-h MR-562 Squamata Mosasauridae Globidens alabamaensis d 0 248 NA-342 $-l-h MR-563 Squamata Mosasauridae Plotosaurus bcnnisoni b/c/f/h 0 248 NA-342 $-l-h MR-564 Squamata Mosasauridae Plotosaurus tuckcri c/i 0 248 NA-343 $-i-h MR-565 Squamata Mosasauridae Platecarpus lympaniticus b/e 0 248 u» Ji. TABLE G.2 Marine Reptile Database Taxa

Loc Map MMR Order Family Genus Species Complete . Reference ID Inierval ID ' vntion

NA-344 $-i-h MR-566 Squaniala Mosasauridae I'lalecarpus coryphaeus b/e 0 248 NA-345 $-i-h MR-567 Squamala Mosasauridae Plalecarpus iciericus b/e/h 0 248 NA-346 $-i-h MR-1225 Plesiosauria Elasmosauridac indel. indel. e 5 270 NA-346 $-i-h MR-568 Squamata Mosasauridae indel. indet. c/e 0 248 NA-347 $-l-h MR-569 Squamala Mosasauridae Eclenosaurus clidasioides b/e 0 248 NA-348 $-i-h MR-570 Squamala Mosasauridae Ectenosaurus clidasloides a 0 248 NA-349 S-i-h MR-571 Squamala Mosasauridae Plioplalecarpus primaevus b/e/h 0 248 NA-35 S-a-c MR-60 Ichlhyosauria indel. indel. indel. c/f/c 3/4 290 NA-350 J-j-h MR-572 Squamala Mosasauridae Plioplalecarpus depressus b/e 0 248 NA-351 $-i-h MR-573 Squamala Mosasauridae Plioplalecarpus crassartus e 0 248; 153 NA-352 j-j-h MR-574 Squamala Mosasauridae Prognalhodon ovenoni b/e/h/i 0 248 NA-353 j-j-h MR-575 Squamala Mosasauridae Prognathodon overtoni L 0 248 NA-354 j-j-h MR-1093 Plesiosauria Elasmosauridae indel. indel. e 4 229 NA-354 j-j-h MR-576 Squamala Mosasauridae Prognalhodon rapax b 0 248 NA-355 $-i-h MR-577 Squamala Mosasauridae Plesioiylosaurus crassidens b/c 0 248 NA-356 j-j-h MR-579 Squamala Mosasauridae ilalisaunis plalyspondylus b/e 0 248 NA-357 $-i-h MR-1473 Squamala Mosasauridae Clidastes liodonlus b/c/d/e 6 M.J. Everhart, personal communicalion 1997 NA-357 $-i-h MR-1476 Squamata Mosasauridae indei. indel. S 0 M.J. Everhan, personal communicalion 1997; 248 NA-357 $-i-h MR-580 Squamala Mosasauridae Tylosaurus prorigor b/e 0 248 NA-358 $-i-h MR-581 Squamala Mosasauridae Tylosaurus nepaeolicus b/e 0 248 NA-359 $-i-h MR-582 Squamala Mosasauridae indel. indel. d/e 0 248; 319 NA-360 $-i-h MR-583 Squamala Mosasauridae Clidasies sp. z 0 248 NA-361 $-$-h MR-584 Squamala Mosasauridae indel. indel. L 0 248 NA-362 $-$-h MR-585 Squamala Mosasauridae Mosasaurus sp d 0 248 NA-363 $-$-h MR-S86 Squamala Mosasauridae Mosasaurus? sp. e 0 248 NA-3M $-i-h MR-587 Squamala Mosasauridae Mosasaurus conodon z 0 248 NA-365 S-i-h MR-1074 Squamala Mosasauridae Clidasies prop>lhon b/e/h/i 5 249 NA-365 $-i-h MR-1090 Squamala Mosasauridae indel. indel. c 5 249 NA-365 $-i-h MR-1073 Squamala Mosasauridae indel. indel. b/c 5 249; 146; 151 NA-365 $-i-h MR-1083 Squamata Mosasauridae Prognalhodon sp. b/c/d/e/i 6 249 NA-365 $-i-h MR-S88 Squamala Mosasauridae Tylosaurus sp L 0 248 NA-366 $-i-h MR-1089 Squamala Mosasauridae indel. indel. e 5 249 NA-366 $-i-h MR-589 Squamata Mosasauridae Plalecarpus sp. z 0 248 NA-366 $-i-h MR-1082 Squamala Mosasauridae Prognathodon sp. b 5 249 NA-366 $-i-h MR-1086 Squamala Mosasauridae Tylosaurus sp. c 5 249 NA-367 $-i-h MR-590 Squamata Mosasauridae Mosasaurus? sp c 0 248 NA-368 $-i-h MR-591 Squamata Mosasauridae Mosasaurus sp d 0 248 NA-369 S-i-h MR-592 Squamala Mosasauridae Globidens? sp. c 0 248 u> TABLE G.2 Marine Reptile Database Taxa

Loc Map MMR Order Tamily Genus Specics Complete . " Reference ID Interval ID • • vntirvn

NA-370 j-j-h MR-593 Squamala Mosasauridae Mosasaurus maximus a 0 248 NA-371 $-i-h MR-595 Squamata Mosasauridae Clidastes sp. b 0 248 NA-371 $-i-h MR-594 Squamala Mosasauridae Tylosaurus sp. b 0 248 NA-372 $-i-h MR-596 Squamala Mosasauridae Plalecarpus sp. b 0 248 NA-373 $-$-h MR-597 Squamala Mosasauridae Platecarpus sp. b 0 248 NA-374 $-i-h MR-598 Squamala Mosasauridae Plalecarpus cf. P. somcnensis z 0 248 NA-375 $-i-h MR-599 Squamala Mosasauridae Clidastes propython b/z 0 248 NA-376 $-i-h MR-600 Squamata Mosasauridae Clidastes propython b/z 0 248 NA-376 $-i-h MR-604 Squamala Mosasauridae Plalecarpus icterius z 0 248 NA-377 $-i-h MR-601 Squamata Mosasauridae Platecarpus icterius b/z 0 248 NA-378 $-i-h MR-602 Squamata Mosasauridae Plalecarpus ictcrius b/z 0 248 NA-379 S-i-h MR-603 Squamata Mosasauridae Platecarpus icterius z 0 248 NA-380 $-i-h MR-605 Squamata Mosasauridae Plalecarpus icterius b/z 0 248 NA-381 $-l-h MR-IOSO Plesiosauria Polycoiylidae Dolichorhynchops osbomi b/c/d/z 6 56 NA-381 $-i-h MR-606 Squamata Mosasauridae Platecarpus icterius b/z 0 248 NA-382 $-i-h MR-607 Squamata Mosasauridae Plalecarpus cf. P. somenensis b/z 0 248 NA-383 $-i-h MR-608 Squamata Mosasauridae Plalecarpus sp. z 0 248 NA-384 S-i-h MR-609 Squamata Mosasauridae Platecarpus sp. b/c/d/h 0 248 NA-385 $-i-h MR-IOSI Plesiosauria Polycoiylidae Dolichorhynchops osbomi b/c/d 6 56 NA-385 $-i-h MR-610 Squamata Mosasauridae Platecarpus sp. z 0 248 NA-386 $-i-h MR-I09I Squamata Mosasauridae Globidens dakoiensis b/d/e 5 250 NA-386 $-l-h MR-6II Squamata Mosasauridae Platecarpus sp. z 0 248 NA-387 S-i-h MR-612 Squamata Mosasauridae Tylosaurus prorigor b/z 0 248 NA-388 S-i-h MR-613 Squamata Mosasauridae Tylosaurus prorigor b/z 0 248 NA-389 S-i-h MR-614 Squamata Mosasauridae Tylosaurus prorigor b/z 0 248 NA-390 j-j-h MR-615 Squamata Mosasauridae Mosasaurus conodon z 0 248 NA-391 j-j-h MR-616 Squamata Mosasauridae Mosasaurus conodon z 0 248 NA-392 j-j-h MR-617 Squamata Mosasauridae Mosasaurus missouriensis b/z 0 248 NA-393 j-j-h MR-618 Squamata Mosasauridae Mosasaurus missouriensis b/z 0 248 NA-394 S-i-h MR-619 Squamata Mosasauridae Clidastes propython b/c/d/e/f/z 0 269;248 NA-42 S-a-b MR-151 Ichthyosauria Mixosauridac Mixosaurus naians e/h/i 5 192;20 NA-42 S-a-b MR-148 Ichthyosauria Shasiasauridae Cymbospondylus petrinus b/c/d/c/f/g/h 6 192;20 /i NA-43 S-d-e MR-75 Ichthyosauria indct. ?Ophlhalmosaurus sp. e 3 68 NA-467 S-i-h MR-744 Squamata Mosasauridae Plolosaurus bennisoni b/c/d/e/f/h 6 50; 248 NA-468 S-i-h MR-745 Squamata Mosasauridae Plolosaurus tuckeri e/r 6 50; 248 NA-469 S-i-h MR-746 Squamata Mosasauridae Plolosaurus sp. e 5 50; 248 NA-470 S-i-h MR-747 Squamala Mosasauridae Plolosaurus sp e 5 50; 248

ON TABLE G.2 Marine Reptile Database Taxa

Loc Map MMR Order Family Genus Species Complete ' Rererence ID Interval ID ' ' vntmn

NA-471 S-i-h MR-748 Squamata Mosasauridae Plotosaurus tuckeri b/c/d/e/f/h/i 6 50; 248 NA-472 $-i-h MR-749 Squamata Mosasauridae Plesiotylosaurus crossidens b/c/d/e/h 5/6 50 NA-493 S-i-h MR-818 Squamata Mosasauridae ?Clidastes sp. b/cyd/e/f/h/i 6 33 NA-494 $-i-h MR-819 Squamata Mosasauridae Clidastes sp. b/c/d/e 0 298 NA-495 $-i-h MR-820 Squamata Mosasauridae Mosasaurus ?missouricnsis b/c/d/e 4/5 298 NA-496 $-i-h MR-821 Squamata Mosasauridae Cilobidens alabamaensis d 2 298 NA-497 $-i-h MR-822 Squamata Mosasauridae Globidens alabamaensis d 2 298 NA-498 S-i-h MR-823 Squamata Mosasauridae Globidens alabamaensis d 2 298 NA-499 S-i-h MR-824 Squamata Mosasauridae Platecarpus coryphaeus b/c/d 0 298 NA-5 j-j-h MR-5 Plesiosauria indet. indet. indet. z 0 233 NA-500 S-i-h MR-825 Squamata Mosasauridae Platecarpus P. somenensis b/c/d 0 298 NA-501 S-i-h MR-826 Squamata Mosasauridae 1'ylosaurus cf. T. prorigor b 0 298 NA-502 j-j-h MR-829 Squamata Mosasauridae ?Plioplatecarpus sp. d 0 298 NA-502 j-j-h MR-828 Squamata Mosasauridae '.'Prognathodon sp. d 0 298 NA-502 j-j-h MR-827 Squamata Mosasauridae '.'Tylosaurus sp. d 0 298 NA-503 S-h-g MR-830 Ichthyosauria indet. indet. indet. e 0 277 NA-504 S-h-g MR-831 Ichthyosauria indet. indet. indet. e 0 277 NA-505 S-h-g MR-832 Ichthyosauria indet. indet. indet. e 0 277 NA-506 S-h-g MR-833 Ichthyosauria indet. indet. indet. e 0 277 NA-510 S-h-g MR-841 Ichthyosauria indet. indet. indet. e 4 194 NA-511 b-b-c MR-842 Ichthyosauria Shastasauridae Shastasaurus sp e/f 6 217; 48; 182 NA-517 S-c-f MR-854 Plesiosauria Elasmosauridae Muraenosaurus Icedsii b/e/f 5 309 NA-517 s-e-r MR-1312 Plesiosauria indet. indet. indet. b/c/d/e/i 5 130 NA-52 S-h-g MR-112 Ichthyosauria Platypterygidae Platypterygius americanus e/f/g/h/i 6 202;17 NA-523 S-i-h MR-860 Squamata Mosasauridae Platecarpus sp. z 5 16 NA-525 S-S-h MR-862 Plesiosauria Polycotylidae Trinacromerum kirki e/f/g/h/i 6 16 NA-526 S-i-h MR-863 Plesiosauria Polycotylidae Trinacromerum kirki e 5 16 NA-529 S-i-h MR-912 Plesiosauria Polycotylidae Dolichorhynchops sp. a 6 309;56 NA-531 S-i-h MR-914 Plesiosauria indet. indet. indet. e/h 0 309 NA-578 S-h-h MR-1047 Plesiosauria PItosauridae Brachauchenius lucasi b 6 56; Bell, personal communication 1999 NA-578 S-h-h MR-1053 Plesiosauria Polycotylidae Trinacromerum bentonianum e/h/i 5 56 NA-579 h-h-h MR-1048 Plesiosauria Pliosauridae Plesiopleurodon wcllesi b/c/d/c/h 6 56; Carpenter, personal communication 1999 NA-580 S-i-h MR-1049 Plesiosauria Pliosauridae Brachauchenius lucasi b/c/d/e/i 6 56; Bell, personal communication 1999 NA-581 S-h-h MR-1052 Plesiosauria Polycotylidae Trinacromerum bentonianum b/c/d/e 6 56 NA-583 S-i-h MR-1056 Plesiosauria Pliosauroidea indet. indet. c 5 253; 309 NA-589 S-i-h MR-1064 Squamata Mosasauridae Clidastes sp. c/d/e 4 268 NA-589 S-i-h MR-1063 Squamata Mosasauridae indet. indet. b/e 4 268 NA-590 j-j-h MR-1065 Squamata Mosasauridae Plioplatecarpus indet. a 6 144 TABLE G.2 Marine Reptile Database Taxa

Loc Map MMR Order I'amily Genus Spccics Complete .. ' Reference ID Interval ID • valion

NA-591 b-b-e MR-1067 Ichthyosauria indet. indet. indet. $ 0 283 NA-594 $-h-g MR-1072 Ichthyosauria Platypterygidae Platypterygius americanus b/c/d 6 242; 173 NA-595 $-i-h MR-1077 Squamata Mosasauridae Clidastes propython e 5 249 NA-595 S-l-h MR-1088 Squamaia Mosasauridae indet. indet. e 5 249 NA-596 $-i-h MR-1303 Squamata Mosasauridae Clidastes liodontus b/c/e/f 5 322 NA-596 $-l-h MR-1078 Squamata Mosasauridae Clidastes propyihon h 5 249 NA-597 $-i-h MR-1079 Squamata Mosasauridae Clidastes propython b/e 5 249 NA-598 $-i-h MR-1092 Plesiosauria Polycotylidae indet. indet. e 4 229 NA-599 $-$-h MR-1094 Pleslosauria Polycotylidae indet. indet. e 4 229 NA-6 j-j-h MR-6 Plesio&auria indet. indet. indet. I 0 233 NA-6 j-j-h MR-7 Squamaia Mosasauridae indet. indet. z 0 233 NA-600 $-a-c MR-1095 Ichthyosauria Shastasauridae Shastasaurus pacificus b/c/d/e/f/i 5 49;182 NA-600 $-8-C MR-1402 Ichthyosauria Shastasauridae loretocnemus califomicus c 5 155 NA-601 $-a-a MR-1097 Ichthyosauria Mixosauridae Mixosaurus cf. M. b/c/d/e/f 5 47 nurdenskioeldii NA-602 $-a-b MR-1098 Ichthyosauria incenae sedis Pessosaurus sp. e/f 6 47 NA-602 $-a-b MR-llOO Ichthyosauria inccrtae sedis Phalarodon cf. P. fraasi b/c/d 5 47; 20 NA-602 $-a-b MR-1099 Ichthyosauria Mixosauridae Mixosaurus cf. M. e/h/i 5/6 47 nordenskioeldii NA-62 $-a-b MR-1325 Ichthyosauria incertae sedis Phalarodon fraasi b/c/d 5 193 NA-62 S-a-b MR-130 Ichthyosauria Mixosauridae Mixosaurus cf M. natans c/f 4 257 NA-62 $-a-b MR-127 Ichthyosauria Mixosauridae Mixosaurus sp. I 3 259 NA-62 $-a-b MR-1396 Plesiosauria Pistosauridae Augustasaurus hagdomi e/f/g/h/1 7 260 NA-629 $-$-h MR-1133 Plesiosauria Elasmosauridae indet. indet. e 5/6 158 NA-63 $-a-b MR-129 Ichthyosauria Mixosauridae Mixosaurus sp. e 4 259 NA-630 S-h-h MR-1134 Plesiosauria Polycotylidae Trinacromerum sp. q/Z 5/6 154;56 NA-664 i-i-h MR-1203 Squamata Mosasauridae cf. Tylosaurus sp. c/f 7 156 NA-670 S-l-h MR-1224 Plesiosauria l-lasmosauridae indet. indet. e 5 270 NA-671 S-i-h MR-1226 Plesiosauria '.'Pliosauroidea indet. indet. e 5 309; 270 NA-672 j-j-h MR-1227 Plesiosauria indet. indet. indet. i 5 270 NA-675 S-d-e MR-1232 Ichthyosauria Ichthyosauridae Baptanodon sp b/c/d/e/f/h 6 89 NA-676 $-f-g MR-1234 Plesiosauria Pliosauridae alT. Pliosaurus sp. b/c/d/e 5 96 NA-682 $-$-r MR-1253 Ichthyosauria indet. indet. indet. b/c/d 4 51; 176 NA-683 $-$-f MR-1254 Ichthyosauria indet. indet. indet. b/c/d 4 51; 176 NA-684 $-i-h MR-1255 Plesiosauria HIasmosauridae Alzadasaurus? sp. nimii 6 77 NA-685 $-i-h MR-1256 Squamata Mosasauridae Prognathodon cf P. overtoni b/c/d/e/f/i/z 6 140 NA-686 $-c-d MR-1257 Ichthyosauria Ichthyosauridae Ichthyosaurus sp. b/c/d/e/f/i 7 86 NA-692 $-a-b MR-1307 Ichthyosauria incertae sedis Paninatator wapitiensis b/c/d/i 6 211 u>

00 TABLE G.2 Marine Reptile Database Taxa

I.oc Map MMR Order Family Uenus Specics Complcic .. ' Rererence ID Interval ID * * vntmn

NA-693 h-h-h MR-14S6 Plesiosauria Polycoiylidae I'rinacromerum benlonianum a 7 297 NA-693 h-h-h MR-1054 Plesiosauria Polycoiylidae I'rinacromerum benlonianum a 7 297;56 NA-694 c-e-f MR-1268 Plesiosauria indel. indel. indel. e 4 308; 229 NA-695 $-i-h MR-1269 Plesiosauria Pliosauridae Polyplychodon hudsoni b/c/d/e 5 311 NA-696 j-j-h MR-1270 Plesiosauria ?Polycoiylidac indel. indei. h 4 223;10 NA-696 j-j-h MR-1279 Squamala Mosasauridae Ilalisaurus plaiyspondylus b/e 4 11 NA-696 j-j-h MR-1280 Squamala Mosasauridae Prognalhudon rapax b/d/e 4 11 NA-697 S-i-h MR-1271 Plesiosauria indel. indel. indel. h 4 10 NA-698 j-j-h MR-1573 Plesiosauria Pliosauridae indel. indel. $ 0 309; 102 NA-698 j-j-h MR-1272 Squaniala Mosasauridae Ilalisaurus plaiyspondylus e 4 45 NA-698 j-j-h MR-1574 Squamala Mosasauridae Mosasaurus sp. S 0 102 NA-699 j-j-h MR-1273 Plesiosauria ?Pliosauroidea indel. indel. d 2 309; 12 NA-699 j-j-h MR-1274 Squamala Mosasauridae Halisaurus plaiyspondylus e 4 12 NA-699 j-j-h MR-1277 Squamala Mosasauridae Mosasaurus cf. M. conodon d 2 12 NA-699 j-j-h MR-1278 Squamala Mosasauridae Mosasaurus cf. M. dekayi d 2 12 NA-699 j-j-h MR-1276 Squamala Mosasauridae Mosasaurus maximus d/e 4 12 NA-699 J-j-h MR-1275 Squamala Mosasauridae Prognalhodon rapax b 4 12 NA-7 j-j-h MR-8 Plesiosauria indel. indel. indel. t 0 233 NA-700 S-i-h MR-1283 Squamala Mosasauridae Halisaurus sp. d 2 266 NA-700 $-i-h MR-1284 Squamala Mosasauridae indel. indel. d 2 266 NA-700 $-i-h MR-1281 Squamala Mosasauridae Plalecarpus sp. d 2 266 NA-700 $-i-h MR-1282 Squamala Mosasauridae Prognalhodon sp. d 2 266 NA-701 j-j-h MR-1285 Squamala Mosasauridae Mosasaurus maximus a 6 43 NA-71 e-e-f MR-138 Ichlhyosauria indel. indel. indel. c/d 3 53 NA-710 $-i-h MR-1306 Squamala Mosasauridae Plioplaiecarpus primaevus c/d/e/h 5/6 300 NA-7 II $-$-h MR-1308 Plesiosauria indel. indel. indel. e 6 58 NA-712 $-i-h MR-1310 Squamala Mosasauridae indel. indel. e 4 195 NA-712 $-i-h MR-1309 Squamala Mosasauridae Tylosaurus sp d/e 4 195 NA-714 $-a-b MR-1315 Ichlhyosauria Mixosauridae Mixosaurus sp. e/f 0 252 NA-716 5-B-g MR-1317 Plesiosauria l:lasmosauridae indel. indel. e/h 4 301; 252 NA-719 $-c-f MR-1326 Plesiosauria Pliosauridae Megalneusaums sp. i 0 161;36 NA-73 $-a-c MR-142 Ichlhyosauria Shaslasauridae Calilbmosaurus peirini b/e/f/h/i 6 192; 48; 20 NA-73 $-a-c MR-141 Ichlhyosauria Shaslasauridae Merriamia /.illeli b/e/r/g/h/i 6 192; 20 NA-73 $-a-c MR-144 Ichlhyosauria Shaslasauridae Shaslasaurus paciflcus b/e/f/h/z 5/6 192; 182;20 NA-73 $-a-t MR-145 Ichlhyosauria Shaslasauridae Shaslasaunis paciricus e/f/h 6 192; 182;20 NA-73 $-a-c MR-146 Ichlhyosauria Shaslasauridae Shasiasaunis paciflcus e/f/i/z 4/6 192; 182; 20 NA-73 $-a-c MR-147 Ichlhyosauria Shaslasauridae Shaslasaurus pacificus e/r/i 5/6 192; 182;20 NA-73 $-a-e MR-143 Ichlhyosauria Shaslasauridae Shaslasaurus pacillcus b/c/e/f/h/i/z 5/6 192; 182;20 U> •ji. so TABLE G.2 Marine Reptile Database Taxa

l.oc Map MMR Order Family Genus Spccies Cumpleic " Rererence ID 1Interval ID * ' vntir^n

NA-73 $-a-c MR-140 Ichthyosauria Shasiasauridae Toretocnemus caliromicus b/e/r/h/i 6 192; Callaway in 254 NA-734 $-$-c MR-1348 Ichthyosauria indel. indet. indel. $ 0 48 NA-736 b-b-c MR-1350 Ichthyosauria indel. indel. indel. h/c/d 0 48 NA-737 b-b-c MR-I3SI Ichthyosauria indel. indet. indel. S 0 48 NA-74 $-a-b MR-150 Ichthyosauria Shasiasauridae Cymbospondylus nevadanus dtlili 5/6 192; 20 NA-74 $-a-b MR-149 Ichthyosauria Shasiasauridae Cymbospondylus piscosiis e 6 192; 20 NA-75 $-h-g MR-152 Ichthyosauria indel. indet. indel. c 2 191 NA-757 $-a-a MR-1393 Ichthyosauria Shasiasauridae Cymbospondylus sp. e/i/z 4 164 NA-8 j-j-h MR-9 Plesiosauria indet. indet. indel. z 0 233 NA-817 S-g-B MR-I3I8 Plesiosauria Polycolylidae indet. indel. e/f/i 0 301; 252 NA-818 i-i-h MR-1468 Plesiosauria Elasmosauridae indet. indel. z 0 309; 251 NA-819 $-i-h MR-1469 Squamata Mosasauridae Plalecarpus sp. i 0 248 NA-82 $-c-d MR-I6I Plesiosauria Plesiosauridae indet. indel. e/i 5 205 NA-820 S-i-h MR-1474 Squamala Mosasauridae Globidens sp. c/d 5 M.J. Hverhart, personal communicalion 1997 NA-821 $-l-h MR-1477 Squamata Mosasauridae indet. indel. $ 0 M.J. Bvethart, personal communicalion 1997; 248 NA-822 $-i-h MR-1478 Squamata Mosasauridae Tylosaurus nepaeolicus $ 0 M.J. Everhart, personal communicalion 1997 NA-823 S-i-h MR-1479 Squamata Mosasauridae indel. indel. S 0 M.J. Everhart, personal communication 1997; 248 NA-824 $-l-h MR-1480 Squamata Mosasauridae indel. indet. S 0 M.J. Everhart, personal communicalion 1997; 248 NA-824 $-i-h MR-I48I Squamata Mosasauridae Tylosaurus nepaeolicus $ 0 M.J. Everhart, personal communicalion 1997 NA-824 S-i-h MR-1482 Squamata Mosasauridae Tylosaurus sp. $ 0 M.J. Everhart, personal communication 1997 NA-850 j-j-h MR-1527 Squamata Mosasauridae Mosasaurus cf. M. missouriensis b/d/e 0 299 NA-850 j-j-h MR-1528 Squamata Mosasauridae Plalecarpus cf. P. somenensis b/c/d/e/h 0 299 NA-851 $-i-h MR-1529 Squamala Mosasauridae Tylosaurus prorigor b/d 0 299 NA-852 S-S-e MR-1532 Ichthyosauria Ichthyosauridae Baplanodon sp. b/c/d/c/f/h/i 0 112 NA-853 $-$-e MR-1533 Ichthyosauria Ichthyosauridae Baptanodon sp. e/h 0 112 NA-854 $-i-h MR-1534 Squamata Mosasauridae Clidasies sp. e 6 140 NA-855 S-i-h MR-1535 Squamata Mosasauridae Mosasaurus sp. z 0 140 NA-859 S-i-h MR-I54I Squamata Mosasauridae Tylosaurus sp. $ 0 14 NA-860 S-i-h MR-1543 Squamata Mosasauridae Mosasaurus dekayi d 0 14 NA-860 S-i-h MR-1542 Squantata Mosasauridae Mosasaurus ma.\imus d 0 14 NA-864 h-h-h MR-1547 Plesiosauria indel. indel. indel. e 6 319 NA-865 S-i-h MR-IS48 Plesiosauria Elasmosauridae indet. indel. $ 0 251 NA-865 S-i-h MR-1549 Squamala Mosasauridae cf. Plalecarpus sp $ 0 251 NA-866 S-i-h MR-I55I Plesiosauria indet. indet. indel. $ 0 157; 251 NA-866 S-i-h MR-1550 Plesiosauria Polycolylidae indel. indel. S 0 157; 251 NA-867 S-S-h MR-1552 Plesiosauria Elasmosauridae indet. indet. e 0 157 NA-868 S-S-h MR-1553 Squamala Mosasauridae indet. indel. e 6 157 NA-869 S-S-h MR-1554 Squamala Mosasauridae indet. indel S 0 157 OJ O TABLE G.2 Marine Reptile Database Taxa

Loc Map MMR Order Family Genus Species Complete . " Rerercnce ID Immal ID VAliiin

NA-870 $-i-h MR-1555 Squamala Mosasauridae indet. indet. S 0 157 NA-872 $-i-h MR-1557 Squamala Mosasauridae indct. indet. e 0 278 NA-873 $-i-h MR-1558 Squamala Mosasauridae indet. indel. c 0 278 NA-874 c-c-r MR-1559 Plesiosauria Polycotylidae indet. indct. c/i 0 301 NA-875 c-c-f MR-IS60 Ichlhyosauria Ichthyosauridac alT. sp. i 0 301 Ophthalmosaurus NA-876 $-e-f MR-I56I Ichthyosauria Ichthyosauridae Ophthalmosaurus sp. b/c 0 301 NA-878 $-i-h MR-1563 Plesiosauria Pliosauridae indct. indct. z 0 307; 309; 251 NA-879 S-i-h MR-1564 Plesiosauria Pliosauridae indet. indet. i 0 307; 309; 251 NA-881 $-i-h MR-IS66 Squamala Mosasauridae indct. indct. $ 0 251 NA-882 $-h-g MR-1567 Ichthyosauria Platyplerygidac Platyplcrygius amcricanus b 4/5 202; 251 NA-883 h-h-h MR-1568 Plesiosauria Polycotylidae indct. indet. i 0 101; 251 NA-884 $-i-h MR-1569 Squamata Mosasauridae Clidastcs? sp $ 0 101 NA-884 $-i-h MR-I570 Squamata Mosasauridae Musasaurus sp $ 0 101 NA-885 $-i-h MR-1571 Plesiosauria Elasmosauridae indet. indet. S 0 102 NA-88S $-i-h MR-! 572 Squamala Mosasauridae cf. Olobidcns sp $ 0 102 NA-886 j-j-h MR-1575 Squamata Mosasauridae Mosasaurus maximus S 0 102 NA-886 j-j-h MR-1576 Squamata Mosasauridae Mosasaurus sp. s 0 102 NA-894 b-b-c MR-1605 Ichthyosauria indct. indct. indct. z 0 145 NA-898 $-i-h MR-I6I0 Squamala Mosasauridae indct. indet. b/c/d 3 124 NA-9 $-i-h MR-II Plesiosauria Polycotylidae Dolichorhynchops obomi z 0 233;56 NA-9 $-i-h MR-IO Squamala Mosasauridae Plaiccarpus ictericus z 0 233 SA-28 e-c-f MR-48 Ichthyosauria Ichthyosauridae Caypullisaurus bonapartci b/c/e/f/h/i 6/7 94; 106 SA-28 e-e-C MR-38 Ichthyosauria Ichthyosauridac Ophthalmosaums sp S 0 106 SA-28 e-c-f MR-39 Plesiosauria Pliosauridae Liopleurodon L. macromerus b/c 0 106 SA-280 $-h-h MR-455 Plesiosauria indct. indet. indel. c/li/i 0 229; 107 SA-281 j-j-h MR-I5I8 Plesiosauria Ulasmosauridac Arisionecles sp. b/c 0 107 SA-281 j-j-h MR-456 Plesiosauria 1-lasmosauridae indct. indel. h/i 0 229 SA-281 j-j-h MR-458 Plesiosauria indel. indet. indct. c/l 0 309;229 SA-281 j-j-h MR-459 Plesiosauria indet. indet. indet. c 0 229; 107 SA-282 j-j-h MR-457 Plesiosauria lilasmosauridae Aristonectes parvidcns a/c/c/i 0 229 SA-283 j-j-h MR-I5I9 Plesiosauria indet. indct. indet. e 0 107 SA-283 j-j-h MR-460 Plesiosauria Polycotylidae indct. indet. e 0 229 SA-284 $-$-<1 MR-461 Plesiosauria indet. indet. indet. c 0 229 SA-29 e-c-f MR-43 Ichthyosauria Ichthyosauridae Ophthalmosaurus monocharaclus a 7 106 SA-29 c-c-f MR-42 Ichthyosauria Ichthyosauridae Ophthalmosaurus sp. S 0 106 SA-29 c-c-f MR-47 Plesiosauria indct. indet. indel. d 1 106 SA-30 c-e-f MR-49 Ichthyosauria indct. indet. indel. S 0 106 TABLE G.2 Maiine Reptile Database Taxa l.oc Map MMR Order Family Genus Species Coniplele . * Rcfcrencc ID Interval ID

SA-30 e-c-f MR-50 Plesiosauria Pliosauridae Pliosaurus sp. $ 0 106 SA-31 c-c-f MR-926 Ichthyosauria Ichthyosauridae Ophihalmosaurus icenicus e/f/h/i 6 37; 108; 94 SA-31 e-e-f MR-53 Ichihyosauria Ichlhyosauridae Ophihalmosaurus sp. S 0 106 SA-31 e-e-f MR-925 Ichlhyosauria indel. indel. indel. e/f 0 37; 108 SA-326 $-$-h MR-547 Squamala Mosasauridae indel. indel. dz 3 231 SA-40 $-d-e MR-1465 Ichlhyosauria Ichlhyosauridae air. sp. b/c 6 282 Ophlhalmosaums SA-40 $-d-c MR-72 Ichlhyosauria incertae sedis ChacaicosBurus cayi b/c/i/h/e 6 93 SA-40 $-d-c MR-1467 Plesiosauria Pliosauridae indel. indel. b/c/d/e 5/6 282 SA-44 $-$-c MR-76 Ichlhyosauria indel. indel. indel. d^i/i 3 287 SA-492 c-e-f MR-921 Ichlhyosauria indel. indel. indel. e 0 176; 37 SA-50 $-f-B MR-IIO Ichlhyosauria Plalyplerygidae Plalyplerygius haulhali h/i 6 17 SA-508 i-i-h MR-83S Squaniala Mosasauridae indel. indel. c/d 4 123;19 SA-518 $-$-d MR-1504 Ichlhyosauria indel. indel. indet. e 0 66 SA-SI8 $-$-d MR-855 Plesiosauria Plesiosauridae indel. indel. e 5 309 SA-519 g-g-8 MR-8S6 Plesiosauria Elasmosauridae Alzadasaurus colombiensis a/b 7 309; 119 SA-519 g-g-g MR-1455 Plesiosauria Pliosauridae Kronosaurus boyacensis a 7 122 SA-534 d-d-e MR-920 Ichlhyosauria indel. indel. indel. e/h/z 0 37; 108 SA-536 $-d-c MR-928 Ichlhyosauria indel. indel. indel. b/c/d 5 176; 37 SA-537 j-j-h MR-1607 Plesiosauria Pliosauridae indel. indel. h/eJtIz 5 59 SA-537 j-j-h MR-931 Squamala Mosasauridae Carinodens cf. C. fraasi d 2 298; 37 SA-537 j-j-h MR-930 Squamala Mosasauridae indel. indel. d 2 37; 146 SA-537 j-j-h MR-929 Squamala Mosasauridae Leiodon cf L. anceps d 2 37; 20 SA-585 h-h-h MR-1059 Ichlhyosauria indel. indel. indel. i 5 19 SA-680 d-d-e MR-1250 Plesiosauria Ctyploclididae cf. Ciyptoclidus sp. e/i 4 109 SA-680 d-d-e MR-1248 Plesiosauria Elasmosauridae cf. Muraenosauius sp. e/i 4 109 SA-680 d-d-c MR-1249 Plesiosauria Pliosauridae indet. indet. e 4 109] SA-687 h-h-h MR-1258 Plesiosauria Elasmosauridae indel. indel. e 5 131 SA-718 g-g-g MR-I32I Ichlhyosauria Plalyplerygidae Plalyplerygius indel. e 4 198 SA-721 $-c-d MR-1328 Ichlhyosauria indel. indel. indel. e 4 108 SA-722 e-«-r MR-1329 Ichlhyosauria Ichlhyosauridae Ophihalmosaurus sp. e/h/i 6 108 SA-723 e-e-f MR-1330 Ichlhyosauria indel. indel. indet. de 5 108 SA-724 e-e-f MR-1331 Ichlhyosauria indel. indel. indel. b/g/i 6 108 SA-725 s-r-g MR-1332 Ichlhyosauria indel. indel. indel. ddiz 5 108 SA-726 $-$-g MR-1333 Ichlhyosauria indel. indel. indel. dV\ 4 108 SA-758 S-i-h MR-1394 Squamala Mosasauridae Ilalisaurus sp. e 5 46 SA-77 $-$-d MR-927 Ichlhyosauria indel. indel. indel. e 0 37; 108 SA-77 $-$-d MR-154 Ichlhyosauria indel. indet. indel. e 2 245; 108 u> K) TABLE G.2 Marine Reptile Database Taxa

Loc Map MMR Preser­ Order Family Genus Species Complete Reference ID Interval ID vation

SA-77 S-S-d MR-923 Ichthyosauria indet. indet. indet. c 0 37; 108 SA-77 MR-155 Ichthyosauria indet. indet. indet. c 2 245; 108 SA-78 $-d-c MR-156 Ichthyosauria Ichthyosauridae Ophthalniosaurus sp. e 2 245; 108; 94 SA-816 $-l-h MR-1464 Squamata Mosasauridae Yaguarasaunis Columbian us hiddldf 6 222 .SA-831 $-e-f MR-1497 Ichthyosauria indet. indet. indet. S 0 104 SA-832 $-c-d MR-1499 Ichthyosauria indet. indet. indet. e 0 66 SA-833 $-c-d MR-1500 Plesiosauria indet. indet. indet. i 0 66 SA-834 $-c-d MR-1502 ichthyosauria indet. indet. indet. e 0 66 SA-835 $-c-d MR-1503 Ichthyosauria indet. indet. indet. e 0 66 SA-836 $-d-e MR-1505 Ichthyosauria indet. indet. indet. e 0 66 SA-837 $-d-c MR-1507 Ichthyosauria indet. indet. indet. z 0 66 SA-838 d-d-e MR-1509 Ichthyosauria indet. indet. indet. b/c/d 0 66 SA-839 d-d-e MR-I5I2 Ichthyosauria indet. indet. indet. c/d/f/i 0 66 SA-839 d-d-c MR-I5I3 Plesiosauria indet. indet. indet. z 0 66 SA-840 $-c-f MR-IS14 Ichthyosauria indet. indet. indet. z 0 66 SA-840 S-e-f MR-1515 Plesiosauria indet. indet. indet. z 0 66 SA-843 j-j-h MR-1520 Plesiosauria indet. indet. indet. i 0 107 SA-844 j-j-h MR-1521 Plesiosauria indet. indet. indet. i 0 107 SA-845 j-j-h MR-1522 Plesiosauria indet. indet. indet. z 5 107 SA-846 j-j-h MR-1523 Plesiosauria Elasmosauridae indet. indet. e 0 107 SA-847 j-j-h MR-1524 Plesiosauria Polycotylidae Trinacromerum lafquenianum e/h 6 107 SA-848 j-j-h MR-1525 Plesiosauria indet. indet. indet. S 0 107 SA-871 $-c-d MR-IS56 Ichthyosauria Ichthyosauridae Ichthyosaurus indet $ 0 65 SA-877 $-c-d MR-1562 Ichthyosauria indet. indet. indet. i 0 67 SA-880 $-d-e MR-1565 Ichthyosauria Ichthyosauridae Ichthyosaurus sp. b/e/r 0 135 SA-889 $-$-c MR-I58I Plesiosauria indet. indet. indet. s 0 60 SA-890 j-j-h MR-1582 Plesiosauria indet. indet. indet. s 0 60 SA-891 j-j-h MR-1583 Plesiosauria indet. indet. indet. s 0 60 SA-892 j-j-h MR-1584 Plesiosauria indet. indet. indet. d/e/f/h/i/z 4/5 60 SA-893 j-j-h MR-1585 Plesiosauria indet. indet. indet. i 6 309; 60 ^Denotes a taxon identified in the literature as living in freshwater habitats (5 records total). These records were not used in data analysis.

U) U) 354

APPENDIX H

DATABASE REFERENCES 355

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APPENDIX I

GRIDDED AND RAW DATA FOR REPTILES AND LITHOLOGIES, PLOTTED ON PALEGEOGRAPHIC BASE MAPS 366

Abbreviations and figure captions for Appendix I

Abbreviations: I = Ichthyosauria P = Piesiosauria M = Mosasauridae

Q = chert V = phosphorite Z = glauconite O = organic-rich rock (> 0.5 weight % TOC)

Figure captions (unrestricted and restricted refer to age resolution of data; see text): I.l. Anisian 1.1.a, Reptile occurrences and intersection with upwelling, age-unrestricted 1.1 .b, Upwelling-related lithology occurrences and intersection with upwelling 1.1 .c. Raw point data, Ichthyosauria, unrestricted 1.1 .d. Raw point data, Piesiosauria, unrestricted 1.1 .e, Raw point data, upwelling-related lithologies

1.2. Norian I.2.a, Reptile occurrences and intersection with upwelling, age-restricted I.2.b, Reptile occurrences and intersection with upwelling, age-unrestricted I.2.C, Upwelling-related lithology occurrences and intersection with upwelling I.2.d, Raw point data, Ichthyosauria, unrestricted I.2.e, Raw point data, Piesiosauria, unrestricted 1.2.f. Raw point data, upwelling-related lithologies

1.3. Pliensbachian I.3.a, Reptile occurrences and intersection with upwelling, age-restricted 1.3.b. Reptile occurrences and intersection with upwelling, age-mirestricted 1.3.c, Upwelling-related lithology occurrences and intersection with upwelling I.3.d, Raw point data, Ichthyosauria, unrestricted I.3.e, Raw point data, Piesiosauria, unrestricted 1.3.f. Raw point data, upwelling-related lithologies

1.4. Callovian I.4.a, Reptile occurrences and intersection with upwelling, age-restricted 1.4. b. Reptile occurrences and intersection with upwelling, age-unrestricted L4.C, Upwelling-related lithology occurrences and intersection with upwelling I.4.d, Raw point data, Ichthyosauria, unrestricted I.4.e. Raw point data, Piesiosauria, unrestricted 1.4.f, Raw point data, upwelling-related lithologies

1.5. Kimmeridgian I.5.a, Reptile occurrences and intersection with upwelling, age-restricted I.5.b, Reptile occurrences and intersection with upwelling, age-unrestricted I.5.C, Upwelling-related lithology occurrences and intersection with upwelling 367

I.5.d, Raw point data, Ichthyosauria, unrestricted 1.5.e. Raw point data, Plesiosauria, unrestricted 1.5.f. Raw point data, upwelling-related lithologies

1.6. Valanginian I.6.a, Reptile occurrences and intersection with upweiling, age-restricted I.6.b, Reptile occurrences and intersection with upweiling, age-unrestricted I.6.C, Upwelling-related lithology occurrences and intersection with upweiling I.6.d, Raw point data, Ichthyosauria, uru-estricted I.6.e, Raw point data, Plesiosauria, unrestricted 1.6.f, Raw point data, upwelling-related lithologies

1.7. Aptian I.7.a, Reptile occurrences and intersection with upweiling, age-restricted I.7.b, Reptile occurrences and intersection with upweiling, age-unrestricted I.7.C, Upwelling-related lithology occurrences and intersection with upweiling I.7.d, Raw point data, Ichthyosauria, unrestricted I.7.e, Raw point data, Plesiosauria, unrestricted 1.7.f, Raw point data, upwelling-related lithologies

1.8. Cenomanian I.S.a, Reptile occurrences and intersection with upweiling, age-restricted I.8.b, Reptile occurrences and intersection with upweiling, age-unrestricted I.8.C, Upwelling-related lithology occurrences and intersection with upweiling I.8.d, Raw point data, Ichthyosauria, unrestricted I.8.e, Raw point data, Plesiosauria, uru-estricted 1.8.f, Raw point data, upwelling-related lithologies

1.9. Coniacian I.9.a, Reptile occurrences and intersection with upweiling, age-restricted I.9.b, Reptile occurrences and intersection with upweiling, age-unrestricted I.9.C, Upwelling-related lithology occurrences and intersection with upweiling I.9.d, Raw point data, Plesiosauria, unrestricted I.9.e, Raw point data, Mosasauridae, unrestricted 1.9.f, Raw point data, upwelling-related lithologies

1.10. Maastrichtian I.lO.a, Reptile occurrences and intersection with upweiling, age-restricted I.lO.b, Upwelling-related lithology occurrences and intersection with upweiling I.lO.c, Raw point data, Plesiosauria, unrestricted I.lO.d, Raw point data, Mosasauridae, unrestricted I.lO.e, Raw pomt data, upwelling-related lithologies 368

Reptile Orders unrestricted

^ p ' D Intersects predicted upwelling (5) ol + P (3) Grid cells by lithology

Q (3) • Intersects predicted upwelling (9) • V (3) • Reptile fossil present (unres.) (3) • V+Z (2) • Z (4) 370

1.1.c.

(chthyosauria Unrestricted point data . Incertae sedis (16) family Indet. (27) A Mixosauridae (22) • Shastasauridae (22)

Plesiosauria Unrestricted point data ~ Plesiosauriaindet (3) A Pistosauridae (4) Upwelling-related lithologies Point data • Q (3) aV (6) Z (7) • O Reptile orders Restricted

' (^) • Intersects predicted upwelling (3) • l + P (1)

Reptile orders Unrestricted

-1 (21) • Intersects predicted upwelling (16) • P (3) • l + P (4) Grid cells by lithology • O (2) Q (17) • Q+V (1) • Intersects predicted upwelling (10) El Q+V+0 (1) • Reptile fossil present (unres.) (8) g Q+Z (1) • V (1) • Z (2) 374

l.2.d.

Ichthyosauria • Ichthyosauridae (1) Unrestricted point data incertae sedis (2) ^milyindet. (24) • Leptopterygldae (1) A Mixosauridae (2) • Shastasauridae (17) ^ Shonisauridae (2) l.2.e.

Plesiosauria * Elasmosauridae (1) Unrestricted point data ; incertae sedis (1) family indet (13) A PIstosauridae (1) T Pleslosauridae (1) Upwelling-related lithologies Point data • Q (39) AV (6) Z (5) 376

Reptile orders restricted I (2) • P (1) l.3.b.

Reptile orders unrestricted : I (6) • P (7) • Intersects predicted upwelling (7) • l+P (9) Grid cells by lithology

O (2) • Intersects predicted upwelling (6) Q (6) Q+Z (1) • Reptile fossil present (unres.) (7) V (3) V+Z (1) Z (3) 378 l.3.d.

\"~^A

Ichthyosauria • Ichthyosauridae (13) Unrestricted point data • family indet. (34) •k Leptopterygidae (13) • (2) T Stenopterygidae (38) Temnodontosauridae (14)

l.3.e.

Plesiosauria •, Elasmosauridae (8) Unrestricted point data c incertae sedis (4) - ^mily indet (23) T Plesiosauridae (15) ^Pliosauridae (13) Upwelling-related lithologies Point data + Q (19) AV (8) Z (8) • o Reptile orders restricted

.•2I (o)ie! • Intersects predicted upwelling (4) • 12 (2)

Reptile orders unrestricted

^ p D Intersects predicted upwelling (9) • l + P (5) Grid cells by lithology • O (6) i i Q (13) • Q + V (1) • Q + Z (1) • Intersects predicted upwelling (16) • V (1) • Reptile fossil present (6) • O + V (1) • V + Z (4) • Z (19) 382

Ichthyosauria Unrestricted point data • Ichthyosauridae (10) ; incertaesedis (1) - family indet. (11) T Stenopterygidae (1)

• Cryptoclidldae (9) Plesiosauna • Elasmosauridae (18) Unrestricted point data - family indet (11) T Plesiosauridae (2) • Pliosauridae (21) Pliosauridae? (1) Upwelling-related lithologies Point data • q (30) qv (2) Av (5) • vz (4) z (51)

0 OC Reptile Orders restricted

^ p • Intersects predicted upwelling (6) • l + P (8)

Reptile Orders unrestricted

^ p • Intersects predicted upwelling (9) • l+P (11) Grid cells by llthology • O (3) L! Q (24) _ • 0-»^Q (1) Q Intersects predicted upwelling (17) • Q-fV (2) •Reptile fossil present (14) B Q + Z (1) • V (1) • V + Z (6) • 2 (11) • O + Z (1) 386 l.5.d.

Ichthyosauria Unrestricted point data • Ichthyosauridae (21) family Indet. (55) • Platypterygidae (1)

• ?Cryptoclididae (1) • Cryptoclididae (18) • Eiasmosauridae (1) • family indet (44) T Plesiosauridae (3) • Pliosauridae (57) ^ Polvcotvlidae (1) Upwelling-related lithologies Point data + Q (79) Q + V (1) • V (9) • V + Z (13) Z (30)

• 0 Reptile Orders restricted I (3) • P (1)

•/r / i.i \ \\

Reptile Orders unrestricted -1 (4) _ «P (4) • Intersects predicted upwelling (2) • l + P (1) I.6.C.

V£ '

Grid cells by llthology • O (3) IJ Q (22) ^ g y (3) • Intersects predicted upwelling (14) • V-^Z (5) • Reptile fossil present (6) • O + V + Z (1) • Z (18) • 0 + Z (1) w VO Ichthyosauria Unrestricted point data family indet. (7) • Platypterygidae (2)

Plesiosauna * Elasmosauridae (2) Unrestricted point data • Elasmosauridae? (1) T Plesiosauridae (2) -AT Pliosauridae (4) •k Pliosauroidea (3) Upwelling-related lithologies Point data + Q (38) AV (15) • V + Z (8) Z (51)

# OC 392

rf

Reptile Orders restricted

0 p • Intersects predicted upwelling (3) • l + P (3)

jJ T

—f

Reptile Orders unrestricted 1 (5) • P (13) ^ Intersects predicted upwelling (9) •l+P (5) Grid cells by llthology • O (7) ! : Q (19) • Intersects predicted upwelling (21) B 0 + Q (1) L'J Q + Z (4) • Reptile fossil present (6) • V + Z (3) • 0 + V + Z (1) • z (25) 0 + Z (1) B Ul u>vo 394

\-v

Ichthyosauria Unrestricted point data • family indet. (11) • Platypterygidae (3)

•r

• Elasmosauridae (9) Plesiosauria • family indet. (8) Unrestricted point data T Plesiosauridae (3) T Plesiosauroidea (1) • Pliosauridae (4) •k Pliosauroidea (2) ^ Polycotylidae (4) Upwelling-related lithologies Point data • Q (34) • Q + Z (1) • V (2) • V + Z (5) " Z (80)

• 0 396

Reptile Orders restricted

' • Intersects predicted upwelling (13) • I + P (2)

Reptile Orders unrestricted

n p • Intersects predicted upwelling (22) • l + P (10) I.8.C.

O (17) Grid cells by lithology u Q (23) m O + Q (3) m Q + V (2) Q + V + 2 (3) Q + 2 (5) Q Intersects predicted upwelling (54) V + Z (11) • Reptile fossil present (unres.) (24) m 0 + V + Z (1) • 2 (35) O + Z (2) VO -J Ichthyosauria Unrestricted point data family indet. (20) • Platypterygidae (20)

V

/•r'

Plesiosauria Unrestricted point data • Elasmosauridae (15) - family indeL (13) -A-Pliosauridae (10) ^ Polycotylidae (8) Upwelling-related lithologies • Q (94) Q + V (2) AV (34) • V + Z (34) Z (146) • O 400

Reptile Orders Restricted • P (2) • Intersects predicted upwelling (2) M (2)

Reptile Orders unrestricted

* r. ion! D Intersects predicted upwelling (33) C M (22) • (29) • 0 (8) . Q (23) • O Q (2) • Intersects predicted upwelling (37) • 0 + Q + V (1) 'Reptilefossil present (25)

LI Q + Z (3) • V (6) • V + Z (8) • V + Z + Q (1) • Z (26) • O + Z (2) 402

Plesiosauria •k Plesiosaurja?Pliosauroidea (4) Unrestricted point data • PlesiosauriaCryptoclididae (1) • PleslosauriaElasmosauridae (54) • Plesiosauriaindet. (30) T PleslosauriaPleslosauridae (1) •k PlesiosauriaPllosauridae (18) •k PleslosauriaPllosauroidea (4) ^ PleslosauriaPolycotylidae (29)

Mosasauridae - sub^mlly Indet (51) Unrestricted point data ^ incertae sedis (1) • Mosasaurinae (80) A Plioplatecarpinae (61) •Tylosaurinae (24) 403

Upwelling-related lithologies Point data • Q (55) :Q + V (4) • Q + Z (1) AV (15) • V + Z (13) - Z (79) • o Reptile Orders restricted

* M (16) ^ Intersects predicted upwelling (20) • P + M (17) • O (8)

I Q (27) m o + Q (1) • Q + V (3) ^ . • Q + V + Z (3) O Intersects predicted upwelling (41)

( I Q + Z (5) • Reptile fossil present (19) • O + Q + Z (1) • V (7) • Q V (6) • Z (28) 4^ o O + Z (3) Ul • ?Pliosauroidea (2) Plesiosauria ^ ?Polycotylidae (1) Unrestricted point data • Elasmosauridae (32) • family indet. (31) -k Pllosauridae (2) ^ Polycotylidae (5)

Mosasauridae - subfamily Indet (43) Unrestricted point data > incertae sedis (3) > Mosasaurinae (64) A Plioplatecarplnae (42) • Tylosaunnae (6) Upwelling-related lithologies Point data • Q (65) Q + V (6) • Q + V + Z (2) • Q + Z (4) AV (40) • V + Z (9) Z (131) 408

APPENDIX J

DISTANCE CALCULATION RESULTS 409

centroid

FIGURE J. 1. Schematic showing possible relationships among adjacent grid cells: A) grid cells share one side, B) grid cells share a vertex with one grid cell intervening, C) grid cells share a vertex with two grid cells intervening. Note that gridding the data can separate closely spaced data points (for example, data scattered about the region where the 6 grid cells share a vertex) because distances were calculated from grid cell centroids.

TABLE J. 1. Calculated distances among all closely spaced data occurrences for the Cenomanian map interval. Distances were calculated using Equation 3 (see text). Cases A, B, and C refer to Figure J. 1. above. These calculations were used to determine the number of adjacent lithology and reptile occurrences, shown in Table 3.12. Case A Case B Case C

Range of distances, km 437-528 796-916 925-1055

Mean distance, km 494 856 990

Number of grid cells 138 223 95 410

Anisian Shortest Distances

7.5_

^ 5.0. g s O" u

0.0, 1000 2000 3000 4000 5000 6000

km between a reptile occurrence and a lithology occurrence

Figure J.2. Shortest distances between reptile and lithology occurrences for the Anisian (age-unrestricted data). 411

Norian Shortest Distances (age-res.)

3-, o Cu 3 O" u

1000 2000 3000 4000 5000 6000 7000 km between a reptile occurrence and a lithology occurrence

Norian Shortest Distances (age-unres.)

1000 2000 3000 4000 SOOO 6000 7000 km bet\veen a reptile occurrence and a lithology occurrence

Figure J.3. Shortest distances between reptile and lithology occurrences for the Norian . Note differing axis lengths. 412

Pliensbachian Shortest Distances (age-res.)

3-,

km between a reptile occurrence and a lithology occurrence

Pliensbachian Shortest Distances (age-unres.) I0.0-,

2500 5000 7500 10000 12500

km between a reptile occurrence and a lithology occurrence

Figure J.4. Shortest distances between reptile and lithology occurrences for the Pliensbachian . Note differing axis lengths. 413

Callovian Shortest Distances (age-res.)

co u o*3 u

1000 2000 3000 4000 5000 6000 7000 km between a reptile occurrence and a lithology occurrence

Calbvian Shortest Dstances (age-unres.)

lobo 2000 3000 4000 5000 6000 7000

km between a reptile occurrence and a lithology occurrence

Figure J.5. Shortest distances between reptile and lithology occurrences for the Callovian. Note differing axis lengths. 414

Kimmeridgian Shortest Distances (age-res.)

10.0.,

>. u c 3u cr 5.0. ub.

0.0, 1000 2000 3000 km between a reptile occurrence and a lithology occurrence

Kimmeridgian Shortest Distances (age-unres.)

u.E

1000 2000 3000

km between a reptile occurrence and a lithology occurrence

Figure J.6. Shortest distances between reptile and lithology occurrences for the Kinuneridgian. Note differing axis lengths. 415

Valanginian Shortest Distances (age-res.)

4. .

^ 3;' cu ; |2i:i X I,;

0.00 0.25 0.50 0.75 1.00 km between a reptile occurrence and a lithology occurrence

Valanginian Shortest Distances (age-unres.)

1000 2000 3000 4000

ion between a reptile occurrence and a lithology occurrence

Figure J.7. Shortest distances between reptile and lithology occurrences for the Valanginian. Note differing axis lengths. 416

Aptian Shortest Distances (age-res.)

4.,

3.,

km between a reptile occurrence and a lithology occurrence

Aptian Shortest Distances (age-unres.) T.S-,

S" 5.0 cu

u. 2.5

0.0 1000 2000 3000 4000 5000

km between a reptile occurrence and a lithology occurrence

Figure J.8. Shortest distances between reptile and lithology occurrences for the Aptian. Note differing axis lengths. 417

Cenomanian Shortest Oistances (age-res.) 20,

>>

10: u.

1000 2000 3000 4000 5000

km between a reptile occurrence and a lithology occurrence

Cenotnanian Shortest Distances (age-unres.) 30,

20

u.

1000 2000 3000 4000 5000 km between a reptile occurrence and a lithology occurrence

Figure J.9. Shortest distances between reptile and lithology occurrences for the Cenomanian. Note differing axis lengths. 418

Con^ian Shortest Distances (age-res.)

o-u 0.5

0.0, 1000

km between a reptile occurrence and a lithology occurrence

Conician Shortest Distances (age-unres.)

30-,

1000 2000 3000 4000 5000 km between a reptile occurrence and a lithology occurrence

Figure J.10. Shortest distances between reptile and lithology occurrences for the Coniacian. Note differing axis lengths. 419

M^trichtian Shortest Distances (age-res.)

>• s3

1000 3000 3000 4000 5000

km between a reptile occuircnce and a lithology

Figure J. 11. Shortest distances between reptile and lithology occurrences for the Maastrichtian. Note differing axis lengths. 420

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