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2015-03-16 Quantitative palynological analyses of Albian-Cenomanian (Lower to Upper Cretaceous) strata in the Sverdrup Basin: Insights into paleoecology, paleoclimatology and palynostratigraphy

Sulphur, Kyle

Sulphur, K. (2015). Quantitative palynological analyses of Albian-Cenomanian (Lower to Upper Cretaceous) strata in the Sverdrup Basin: Insights into paleoecology, paleoclimatology and palynostratigraphy (Unpublished master's thesis). University of Calgary, Calgary, AB. doi:10.11575/PRISM/27507 http://hdl.handle.net/11023/2118 master thesis

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Quantitative palynological analyses of Albian-Cenomanian (Lower to Upper Cretaceous) strata

in the Sverdrup Basin: Insights into paleoecology, paleoclimatology and palynostratigraphy

by

Kyle Christopher Sulphur

A THESIS

SUBMITTED TO THE FACULTY OF GRADUATE STUDIES

IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE

DEGREE OF MASTER OF SCIENCE

GRADUATE PROGRAM IN GEOSCIENCE

CALGARY, ALBERTA

MARCH, 2015

© Kyle Christopher Sulphur 2015 Abstract

Multivariate statistical analyses of terrestrial palynomorphs from Albian to Cenomanian (Lower to Upper Cretaceous), Sverdrup Basin strata reveal a landscape with wet lowlands inhabited by

Pteridophytina and Bryophyta, with cooler, moist uplands inhabited by a variety of conifer plants. A humid and temperate climate prevailed at this time. Three sites from eastern Sverdrup

Basin contain similar palynofloras with long-ranging taxa, but quantitative palynological analyses reveal important differences in relative abundances between localities. A total of eleven early angiosperm pollen were recovered, providing further evidence of a delay in the dispersal and diversification of angiosperms into the Sverdrup Basin relative to more southern locations.

This latitudinal diachroneity may potentially be due to paleogeographic barriers, such as, the

Arctic Ocean, the extent of the Western Interior Seaway, or due to limited insect pollination dispersal strategies.

i Acknowledgements

I would first and foremost like to thank my co-supervisors: Drs. Jennifer Galloway (GSC-

Calgary), Federico Krause (University of Calgary), Len Hills (University of Calgary) and committee member Keith Dewing (GSC-Calgary) for their continued dedication and support throughout the writing of this thesis. I would next like to thank Art Sweet for lending his enormous inventory of books, encouraging ideas, and taxonomic support, James White for introducing me to the world of palynology, and Ashton Embry (all three GSC-Calgary) for his great insights into the geology of the Sverdrup Basin. I thank Andrew MacRae (Saint Mary’s

University) for discussing Albian-Cenomanian stratigraphy with me, and along with Rob

Fensome (GSC-Atlantic) for help with the world of dinoflagellate cysts. I thank Len Hills and

Koldo Núñez-Betelu for collecting palynological samples from Strathcona and Cañón fiords and

Claudia Schröder-Adams for palynological sample collection at Glacier Fiord. Thank you to

Linda Dancey for palynological preparation and for curation support. Financial support, lab equipment and office space was contributed by the Geological Survey of Canada-Calgary under the GEM (Geomapping for Energy and Minerals) and Western Arctic project, headed by Keith

Dewing and Jennifer Galloway, to which I am extremely grateful.

And finally, I dedicate this thesis to Len V. Hills, to whom I owe so much.

ii Table of Contents Abstract...... i Acknowledgements...... ii Table of Contents...... iii List of Plates and Tables...... iv

INTRODUCTION ...... 1

REGIONAL CONTEXT ...... 7

STRATIGRAPHY ...... 8 Christopher Formation...... 10 Hassel Formation ...... 11 Bastion Ridge Formation ...... 14 Kanguk Formation ...... 16

METHODS ...... 17 Palynological preparation ...... 19 Palynology ...... 23 Taxonomic Notes...... 23 Multivariate Statistical Analyses ...... 25

RESULTS ...... 26 Angiosperm Pollen...... 26 R-Mode Cluster Analysis...... 29 Multidimensional Scaling Ordination...... 33 CONISS ...... 33 Q-mode Cluster Analysis...... 39

DISCUSSION...... 42 Stratigraphic Palynology...... 42 Angiosperms in Context ...... 43 Angiosperm Biostratigraphy...... 45 Paleoecology and Paleoclimate...... 49 R-Mode Cluster Analysis...... 54 Multidimensional Scaling Ordination...... 56 CONISS ...... 59 Q-Mode Cluster Analysis ...... 61

SUMMARY AND CONCLUSIONS ...... 63

REFERENCES ...... 67

APPENDIX I: Glacier Fiord count data ...... 78 APPENDIX II: Strathcona Fiord count data...... 80 APPENDIX III: Cañón Fiord count data...... 84

iii List of Tables

Table 1: Time constraints by formation...... 9

Table 2: GSC Curation sample numbers ...... 20

Table 3: Taxonomic authorities ...... 24

Table 4: Further taxonomic information...... 51

List of Figures

Figure 1: Map of the Sverdrup Basin and sample locations found in this study ...... 2

Figure 2: Regional Stratigraphy...... 3

Figure 3: Stratigraphy and location of palynomorph samples at Glacier Fiord...... 15

Figure 4: Location of Glacier Fiord section...... 18

Figure 5: Location of Cañón Fiord section ...... 21

Figure 6: Location of Strathcona Fiord...... 22

Figure 7: R-mode Cluster Analysis...... 30

Figure 8: Multidimensional Scaling Ordination ...... 34

Figure 9: Glacier Fiord CONISS ...... 35

Figure 10: Strathcona Fiord CONISS ...... 36

Figure 11: Cañón Fiord CONISS...... 37

Figure 12: Q-Mode Cluster Analysis...... 40

Figure 13: Paleogeographic map of North America during the Albian...... 46

List of Plates

Plate 1...... 27

iv INTRODUCTION

Fossil pollen and spores are abundant in terrestrial and marine rocks and show a large number of distinct morphological characters that facilitate their identification, making them excellent tools for biostratigraphy. In particular, the pollen of angiosperms (the flowering plants) are important to palynostratigraphy in post-Cenomanian (Late Cretaceous) units due to their dominant presence on the landscape. However, the early history of angiosperms is poorly understood, particularly in polar basins such as the Sverdrup Basin (Figure 1).

The Sverdrup Basin is an east-west trending, Carboniferous to Paleogene sedimentary basin that stretches from Melville Island in the west to Ellesmere Island in the east. It is approximately 1,000 km long, 350 km wide and is up to 13 km deep at its depositional center

(Balkwill, 1978). A major Early Cretaceous transgression deposited offshore muds of the

Christopher Formation during the Aptian and Albian. During the late Albian to Cenomanian

(Early to Late Cretaceous), widespread deposition of sand, silt and clay became what is now known as the Hassel Formation. A subsequent Late Cretaceous transgression deposited clay and silt of the Cenomanian to early Maastrichtian Kanguk Formation (Figure 2) (Embry and

Beauchamp, 2008; Schröder-Adams, 2014).

The Albian and Cenomanian ages are a time of early angiosperm expansion across North

America from south to north. For example, the first North American angiosperm pollen are recorded in the early Albian Potomac group of Maryland, USA (Brenner, 1964; Doyle, 1969;

Wolfe et al., 1975; Doyle and Hickey, 1976). Middle Albian angiosperm pollen are next recorded further north in the lower Colorado Group, Swan River Formation and Peace River

1 80° 75° 1). Ellesmere Island A′

Cañón Fiord Jones Sound Sound Jones Strathcona Fiord Devon Island

-90°

Graham -90° Glacier Fiord Glacier Axel Heiberg

Cornwallis Cornwall AmundRingnes

Ellef Bathurst

Ringnes King Christian 400 km

Ocean Ocean Arctic A

Lougheed N

d

Mackenzie King Borden Borden -110° -110° -110° n la Is le il

Brock Brock lv e M LEGEND

Sverdrup Basin (Land) Sverdrup Basin (Water) Sample Locations

Eglinton Eglinton Prince Patrick Patrick Prince M’Clure Strait Banks Island 80° 75° Figure 1: Index map of Canada highlighting the location of the Sverdrup Basin (shaded gray) and Figure 1: Index map of Canada highlighting the location of the Sverdrup from Dewing and Obermajer (201 sampling site locations for this study (black circles). Map modified

2 NE V (Embry, 1991) (Embry, (Embry, 1991) (Embry, SW Ellesmere SW , 1992) , 2014) (Osadetz and Moore, 1994) (Embry, 1991) (Embry, et al. Expedition Fm (Embry and Beauchamp, 2008) (Embry and Beauchamp, Invincible Point Mbr Macdougal Point Mbr (MacRae, 1994) Bastion Ridge Fm Glacier Fiord Glacier Fiord Axel Heiberg, Axel Heiberg, Bastion Ridge Fm (Núñez-Betelu Walker Island Mbr Walker Paterson Island Mbr Rondon Mbr (Schröder-Adams et al. (Schröder-Adams (Embry, 1991) (Embry, , 1983) , 2012) , 2013) et al. et al. et al. Ellef Ringnes, Hoodoo Dome Hoodoo Dome Hassel Fm Hassel Fm (Balkwill Kanguk Fm Isachsen Fm Deer Bay Fm , 2014), 2014) , 2014), al. et al.et (Pugh(Pugh Christopher Fm (Núñez-Betelu, 1994) (Galloway (Galloway , 1982) et al. Lougheed Lougheed Volcanics Sandstone Dominated Mudstone Dominated (Embry and Beauchamp, 2008) V (Balkwill SW Ma 72.1 86.3 93.9 89.8 83.6 ~113 ~133 ~125 ~129 ~140 100.5 Age Aptian Albian Turonian Santonian Coniacian Berriasian

Barremian Campanian Hauterivian Valanginian

Cenomanian Maastrichtian

Location Late Early Early

Epoch Cretaceous Period Figure 2: Generalized Cretaceous chronostratigraphy of the Sverdrup Basin from Southwest (Lougheed Island) to Figure 2: Generalized Cretaceous chronostratigraphy of the Sverdrup A-A′ of Figure 1. Numeric ages for stage boundaries are from ICS (2014). Northeast (Ellesmere Island) following 1 and Galloway et al. (2013) for further chronological control references. Table Refer to

3 Formation of southern Alberta (Norris, 1967; Playford, 1971; Singh, 1971). Late Albian

angiosperm pollen are first documented in the Sverdrup Basin in the upper Christopher and

Hassel formations on Ellef Ringnes Island (Hopkins and Balkwill, 1973; Balkwill and Hopkins,

1976; Galloway et al., 2012). Due to these late Albian angiosperm occurrences on Ellef Ringnes

Island, similar strata elsewhere in the Sverdrup Basin were thought also likely to yield early

angiosperm pollen.

Early angiosperms are thought to have been ecologically competitive in disturbed

lowland settings such as deltas and fluvial environments and are often well preserved in over

bank deposits (Hickey and Doyle, 1977; Retallack and Dilcher, 1986; Herman, 2002; Coiffard et al., 2006; Royer et al., 2010). Therefore, their pollen are most likely to be preserved in rocks reflecting a similar depositional environment. In particular, the coaly mudstones of the Hassel

Formation are interpreted to have been deposited in overbank environments within a floodplain that included riparian and other disturbed lowland environments (Balkwill and Hopkins, 1976;

Balkwill and Roy, 1977; Balkwill et al., 1982; Balkwill, 1983; Galloway et al., 2012). The

Hassel Formation was selected for study to maximize the likelihood of finding early angiosperm

pollen due to its age and depositional environment.

In addition to early angiosperm pollen, Hassel Formation palynological samples provide

numerous examples of other well-preserved pollen and spores representing non-angiosperm

fossil plant groups. To capitalize on the large abundances of fossil pollen and spores recorded in

Hassel Formation sediments, exploratory quantitative multivariate statistics were employed to

examine stratigraphic and ecological relationships. These statistical techniques include: Q- and

4 R-mode hierarchical cluster analyses, multidimensional scaling ordination (MDS) and stratigraphically constrained incremental sum of squares cluster analysis (CONISS).

To delineate ecological groupings irrespective of stratigraphic context, hierarchical cluster analysis were carried out on taxa (R-mode). Cluster analysis is a statistical process used to group sets of variables (in this case taxa) into clusters in such a way that variables in each cluster are more similar to each other than to those in a different cluster. This involves a measurement of distance such as Euclidean distance (in this case representing co-occurrence) and a method of grouping these objects (in this case, Ward’s method). Ward’s method and

Euclidian distance were chosen because these are known for generating meaningful clusters in palynological data (Birks and Gordon, 1985; Kent and Cooker, 1992; Hills and Strong, 2007;

Galloway et al., 2012).

In addition to R-mode hierarchical cluster analysis, multidimensional scaling ordination

(MDS) was performed to independently identify populations of taxa based on palynological composition. Many ordination techniques exist (e.g. principal components analysis (PCA), correspondence analysis (CA), multidimensional scaling (MDS)) and are used to represent multivariate data by reducing the number of variables to two or three principal variables that explain the majority of variation in the data. MDS was chosen over other ordination techniques such as PCA and CA because MDS does not contain assumptions that are present in PCA or CA about species distribution patterns along a compositional gradient (Kruskal, 1964; Prentice,

1977; Minchin, 1987; Clarke, 1993). Ecological groupings as delineated by R-mode hierarchical

5 cluster analysis and MDS can then be interpreted based on comparison with modern

representatives for each taxa grouping to provide the basis for climate inferences.

This thesis also endeavors to refine Early Cretaceous palynostratigraphy. Previous Early

Cretaceous palynological work facilitated age recognition in a broad sense, but the potential for

detailed correlation was limited by the predominance of globally long-ranging fossil types

typical for this time period (Hopkins, 1974; Traverse, 2007; Galloway et al., 2013). This thesis uses the relative abundances of these long-ranging pollen and spores present to examine the stratigraphic potential of Early Cretaceous palynological acme zones based on stratigraphically constrained incremental sum of squares cluster analysis (Grimm, 1987). In addition, this thesis uses hierarchical cluster analysis of samples (Q-mode) to explore relationships between samples based on their palynomorph content and to test the following hypothesis “Samples from Hassel

Formation, irrespective of location (Glacier Fiord, Strathcona Fiord, Cañón Fiord) contain similar palynoassemblages.”

In summary, this thesis investigates the late Albian to Cenomanian palynoflora of the

Hassel Formation and bounding units for multiple purposes: 1. to document early angiosperm pollen in the Sverdrup Basin; 2. employ multivariate statistical analyses to delineate ecological groupings to provide insight into polar late Early and early Late Cretaceous paleoclimate; 3. explore utility of quantitative palynology to better understand palynostratigraphy.

6 REGIONAL CONTEXT

As part of large-scale tectonic plate reorganization during the Early Carboniferous, the

Sverdrup Basin emerged as a rift basin (Embry and Beauchamp, 2008). Extension of the

Sverdrup Basin was intermittent through the Carboniferous and Early Permian, eventually progressing into a period of thermal subsidence that lasted until the Early Cretaceous (Embry and

Beauchamp, 2008). Rift-related subsidence produced a marine basin that initially deposited extensive Upper Carboniferous evaporite sequences. During the Mesozoic, these evaporites came to form diapiric structures within the basin, mostly concentrated on Axel Heiberg, Amund

Ringnes, and Ellef Ringnes islands (Nassichuck and Davies, 1980). Siliciclastic sediments flooded the basin through the rest of the Late Paleozoic and the early Mesozoic, but began to wane by the end of the Jurassic and the earliest Cretaceous (Embry and Beauchamp, 2008).

A widespread unconformity of Hauterivian (Early Cretaceous) age was followed by renewed rifting and extension in the Early Cretaceous, which coincided with seafloor spreading in the adjacent Amerasia Basin (Embry and Beauchamp, 2008). A major Early Cretaceous transgression during the middle Aptian to early Albian drowned the Sverdrup Basin, resulting in accumulation of offshore muds and silts that are now mudstones and siltstones referred to as the

Christopher Formation, which is early to middle Albian in age (Hopkins, 1974; Balkwill and

Hopkins, 1976). Continued sedimentation in the late Albian to Cenomanian resulted in the accumulation of shoreline and shallow shelf deposits of the Hassel Formation (Hopkins and

Balkwill, 1973; Embry and Beauchamp, 2008; Galloway et al., 2012).

7 Thick basaltic flows and dykes/sills were emplaced during the Early Cretaceous, and earliest Late Cretaceous, as exemplified by the Cenomanian Strand Fiord Formation. These igneous rocks are related to a hotspot north of the basin at the time (Embry and Osadetz, 1988;

MacRae 1996; Embry and Beauchamp, 2008; Jowett and Williamson, 2014). A major transgression then allowed deposition of the mudstone and siltstone-dominated Kanguk

Formation, which is Cenomanian to early Maastrichtian in age (Table 1) (Núñez-Betelu, 1994;

Schröder-Adams et al., 2014; Pugh et al., 2014). Sediment input continued and increased towards the end of the Cretaceous, and encompasses the shoreline to shallow marine sandstones of the Expedition Formation. The Sverdrup Basin’s 300 million year history concluded in the

Eocene with widespread uplift associated with the Eurekan Orogen (Embry and Beauchamp,

2008).

STRATIGRAPHY

In the central part of the Sverdrup Basin, the late Albian to early Cenomanian Hassel

Formation conformably lies atop marine mudstones of the Aptian to late Albian Christopher

Formation and is overlain conformably by marine mudstones and claystones of the Cenomanian to Maastrichtian Kanguk Formation (Embry and Dixon, 1990; 1994; Burden and Languille,

1991; Dixon, 1993). This tripartite package is present throughout much of the Canadian Arctic, except on southern and western Axel Heiberg Island, where the Hassel Formation is overlain by the mudstone-dominated Bastion Ridge Formation and the basaltic volcanics of the Strand Fiord

Formation, both of Cenomanian age. Table 1 lists time constraints for these units and Figure 2 shows the regional stratigraphy schematically.

8 Table 1: Summary of time constraints by formation.

Formation Age Based on References Palynomorphs, radiolarians, Fricker, 1963; Wall, 1983; Stott, 1968; Pluchut foraminifera, and macrofossils and Jutard, 1976; Dorenkamp et al., 1976; from Lougheed, Ellef Ringnes, Balkwill and Hopkins, 1976; Miall et al., 1979; Cenomanian to Banks islands, and Eclipse Balkwill et al., 1980; Miall et al., 1980; Balkwill Kanguk Maastrichtian Trough. et al., 1982; Balkwill et al., 1983; Núñez-Betelu et al., 1992; Núñez-Betelu and Hills, 1998; Pugh et al., 2014; Schröder-Adams et al., 2014

Metasequoia Miki (Miki, 1941) Fricker, 1963; Ricketts et al., 1984; Trettin and branchlet impressions on Axel Parrish, 1986; Núñez-Betelu et al., 1992; Heiberg, 40Ar/39Ar dating Tarduno et al., 1998; LePage et al., 2005 indicate an age of 95.3 +/- 0.2 Strand Fiord Cenomanian Ma (near the Cenomanian- Turonian boundary) on Axel Heiberg Island; U/Pb dating 92.0 +/- 1 Ma (Early Turonian) on Ellesmere Island. Palynomorphs, Foraminifera Fricker, 1963; Núñez-Betelu et al., 1992; from Axel Heiberg and Tarduno et al., 1998; Schröder-Adams et al., Bastion Ellesmere Islands, 2014 Cenomanian Ridge Interfingering of radiometrically dated Strand Fiord volcanics (above). Palynomorphs, and Fricker, 1963; Larochelle et al., 1965; Stott, macrofossils from Lougheed, 1968; Hopkins and Balkwill, 1973; Ellef Ringnes and Banks islands, Balkwill and Hopkins, 1976; Pluchut and Jutard, late Albian to and Eclipse Trough 1976; Dorenkamp et al., 1976; Miall et al., 1979; Hassel early Balkwill et al., 1980; Miall et al., 1980; Balkwill Cenomanian et al., 1982; Galloway et al., 2012; Schröder-Adams, 2014

Dinoflagellate cysts, Hopkins and Balkwill, 1973; Balkwill et al., upper palynomorphs from Lougheed, 1982; Galloway et al., 2012 Cenomanian Hassel Ellef Ringnes, and Axel Heiberg islands Macrofossils, palynomorphs, Plauchut and Jutard, 1976; Miall et al., 1979; lower late Albian dinoflagellate cysts from Banks Núñez-Betelu, 1994 Hassel and Ellesmere Islands Palynomorphs, foraminifera, Ficker, 1963; Stott, 1968; Pluchut and Jutard, macrofossils from Lougheed, 1976; Dorenkamp et al., 1976; Miall et al., Mackenzie King, Prince Patrick, 1979; Aptian to Eglinton, Banks, Melville, Axel Balkwill et al., 1980; Balkwill et al., 1982; Christopher Albian Heiberg, Ellesmere and Ellef Balkwill, 1983; Wall, 1983; Harrison, 1995; Ringnes islands. Nøhr-Hansen and McIntyre, 1998; Harrison and Brent, 2005; Hall et al., 2005; Schröder-Adams et al., 2014

9 Christopher Formation

The Christopher Formation was named by Heywood (1955) for a 470 m succession of marine mudstone exposed on Christopher Peninsula, northern Ellef Ringnes Island. It ranges in thickness from 975 m on Amund Ringnes Island, down to 300 m on Banks Island (Plauchut and

Jutard, 1976; Miall et al., 1979; Balkwill, 1983). The Christopher Formation is divided into the lower Invincible Point Member, a mudstone succession with fine to medium-grained sandstones and mudstones, often containing concretions. A 5 to 6 m thick sandstone bed at the top of the

Invincible Point Member is a marker that separates the two members (Balkwill, 1983; Schröder-

Adams et al., 2014). The Invincible Point Member is interpreted as a mid-shelf to lower/upper offshore environment (Embry, 1985). The MacDougall Point Member is interpreted to represent marine shelf conditions (Embry, 1985; Balkwill, 1983; Schröder-Adams et al., 2014).

Macrofossils (Jeletzky fide Plauchut and Jutard, 1976), foraminifera (Fischer fide

Plauchut and Jutard, 1976), palynomorphs (Miall et al., 1979), and dinoflagellates cysts

(Dorenkamp et al., 1976) preserved in Christopher Formation sediments from Banks Island, as well as foraminifera, ostracodes and dinoflagellates cysts from Melville Island (Harrison, 1995) show that in the western part of the Sverdrup Basin the Christopher Formation represents the early to middle Albian. Monocotyledonous angiosperm pollen were recovered from the upper 20 cm of Christopher Formation on Eglinton Island, indicating an extension into the late Albian

(Davies and Wall fide Harrison and Brent, 2005). In the basin center, macrofossils indicating an

Aptian age were found on Amund Ringnes Island (Jeletzky fide Balkwill, 1983). Also on Amund

Ringnes and Ellef Ringnes islands, dicotyledonous angiosperm pollen are present, suggesting that the formation extends to the latest Albian in these locations as well (Hopkins, 1974; Balkwill

10 and Hopkins, 1976). Foraminifera preserved in the Christopher Formation at Glacier Fiord on

Axel Heiberg Island indicate an overall age of late Aptian through late Albian (Schröder-Adams et al., 2014). U-Pb dating of a bentonite (111.74 +/- 0.26 Ma) preserved in the upper Invincible

Point Member at this locality confirms that the succession ranges into the Albian (Herrle et al.,

2014; Schröder-Adams et al., 2014). Therefore, the Christopher Formation is taken to represent

Aptian through latest Albian time in central Sverdrup Basin, with only early to late Albian time preserved in western Sverdrup Basin.

Hassel Formation

The Hassel Formation was named by Heywood (1955 and 1957) for a 550 m succession of sandstone lying on central and eastern Ellef Ringnes Island. Hassel Formation ranges from

450 m on Amund Ringnes Island (Balkwill, 1983), down to 20-50 m on Banks Island (Miall et al., 1979). The Hassel Formation is conformable with the preceding Christopher Formation across the Sverdrup Basin from Melville Island in the west to Ellesmere Island in the east

(Balkwill and Roy, 1977; Balkwill et al., 1982; Balkwill, 1983; Núñez-Betelu, 1994; Harrison,

1995; MacRae, 1996; Galloway et al., 2012; Schröder-Adams et al., 2014).

Two informal members of the Hassel Formation are recognized in the Sverdrup Basin.

The lower member is fine- to medium- grained, poorly lithified sands with thin, low angle cross beds. The upper member contains medium to large scale, high angle cross beds and contains thin carbonaceous mudstone and coal beds (Galloway et al., 2012). The thickness of the Hassel

Formation and its informal members fluctuates throughout the basin due to variable accommodation space associated diapiric intrusion and erosion (Plauchut and Jutard, 1976;

11 Balkwill and Roy, 1977; Balkwill et al., 1982; Balkwill, 1983; Harrison, 1995; Galloway et al.,

2012).

At Hoodoo Dome on Ellef Ringnes Island and on Lougheed Island, a thin bed of ferruginous siltstone appears at the base of the upper member and may represent a divide between the two informal members (Hopkins and Balkwill, 1973; Balkwill and Roy, 1977;

Balkwill et al. 1982; Balkwill, 1983; Galloway et al., 2012). This division could also be an unconformable surface, because on Melville, Axel Heiberg, Ellesmere and Banks islands, the upper member of the Hassel Formation is entirely absent or is not interpreted to be of

Cenomanian age due to a paucity of Cenomanian-indicating angiosperm pollen (Plauchut and

Jutard, 1976; MacRae, 1996; Núñez-Betelu, 1994, Harrison, 1995). This will be discussed further in the Angiosperm Biostratigraphy section. Where this unconformity exists, it is thought to span the late Albian to Cenomanian and thought to represent a first-order sequence boundary associated with the cessation of sea-floor spreading of the Amerasia Basin to the northwest

(Embry and Dixon, 1990; Dixon, 1993; Embry and Dixon, 1994).

The lower member of the Hassel Formation represents a shallowing-upward sequence interpreted to represent a beach or other shore face deposit peripheral to a deltaic or fluvial system (Balkwill and Hopkins, 1976; Balkwill and Roy, 1977; Balkwill et al., 1982; Balkwill,

1983). The upper Hassel Formation is interpreted to be of fluvial (possibly braided) or marginal marine origin on the basis of granule lenses possibly being channel deposits, and high angle cross stratified sandstones as possible point bar deposits. The thin coals and mudstones could be

12 overbank muds and back-swamp deposits (Balkwill and Hopkins, 1976; Balkwill and Roy, 1977;

Balkwill et al. 1982; Balkwill. 1983; Galloway et al., 2012; Pugh et al., 2014).

Evidence for a late Albian to Cenomanian age for the Hassel Formation comes primarily from fossil evidence and volcanic radiometric dates. Paleontological studies of lower Hassel

Formation indicate a late Albian age, as is indicated by ammonites and dinoflagellates cysts

(Jeletzky in Plauchut and Jutard, 1976), pelecypods, gastropods (Jeletzky, 1974 in Miall et al.,

1979) and palynomorphs (Hopkins, Sweet and Brideaux in Miall et al., 1979), all of which were collected from various sites on Banks Island. Late Albian palynomorphs and dinoflagellates cysts from Hassel Formation on Ellesmere Island were also documented by Núñez-Betelu

(1994). Additional paleontological evidence from the upper Hassel Formation also indicates a

Cenomanian age, as Cenomanian to Turonian dinoflagellates cysts were observed in material collected from Lougheed Island (Brideaux in Balkwill et al., 1982). Palynomorphs diagnostic of late Albian to Cenomanian age were collected from Ellef Ringnes Island and late Albian dinoflagellate cysts were recovered from Axel Heiberg Island (Hopkins and Balkwill, 1973;

MacRae, 1996; Galloway et al., 2012).

Notably, radiometric dates from volcanic units corroborate paleontological data, as diabase sills and dykes from the middle of the Hassel Formation on Ellef Ringnes Island yielded

K/Ar radiometric dates 102-110 Ma of Albian age (Larochelle et al., 1965). Upper Hassel sediments that are interbedded with Bastion Ridge Formation flood volcanics from Axel Heiberg

Island were dated as Cenomanian in age using Ar40/Ar39 as 95.3 +/- 0.2 Ma old (Embry and

Osadetz, 1988; Trettin and Parish, 1987; Tarduno et al., 1998; Embry and Beauchamp, 2008).

13 Thus, the age of the Hassel Formation across the Sverdrup Basin ranges from late Albian to

Cenomanian.

In particular, at the Glacier Fiord locality on Axel Heiberg Island, only the late Albian portion of the Hassel Formation is present. Foraminifera from the Miliammina manitobensis/Reophax Zone were collected from this locality (Schröder-Adams et al., 2014).

This zone is indicative of the late Albian stage, based on comparison to the Western Interior

Seaway (Caldwell et al., 1978). Previous palynological studies of Hassel Formation at Glacier

Fiord report palynomorphs that are late Albian in age and are comparable to those recovered from Hassel Formation exposed on Ellef Ringnes Island (Hopkins and Balkwill, 1973; Núñez-

Betelu, 1994; MacRae, 1996; Galloway et al., 2012). A paleosol is recorded at the top of the

Hassel Formation at the Glacier Fiord locality and is immediately followed by the Bastion Ridge

Formation (Figure 3) (Schröder-Adams et al., 2014).

Bastion Ridge Formation

The Bastion Ridge Formation was named by Fricker (1963) for organic-rich mudstone lying atop the Hassel Formation, but below the Kanguk Formation at Bastion Ridge, Axel

Heiberg Island. On Axel Heiberg Island, the only place where the Bastion Ridge Formation is recognized, this unit unconformably overlies the Hassel Formation and is unconformably overlain by the Kanguk Formation (Núñez-Betelu et al., 1994; Schröder-Adams et al., 2014).

MacRae (1996) interpreted the majority of the Bastion Ridge Formation to be deposited in an offshore environment, possibly with dysaerobic bottom conditions. Foraminiferal biostratigraphy of the Bastion Ridge Formation at Glacier Fiord indicate a Cenomanian age based on a

14 Glacier Fiord, southern Axel Heiberg Island

Formation Palynology Meters Grain size: Clay Silt VF F M C Stage Formation samples

Mud sized Sand sized ySstem

Series 380

370 360 Depositional 350 Environment 340 Offshore Turonian 330

Kanguk 320 Bastion Ridge 310

300

93.9 Ma p18-p20 290 Paleosol p16-p17

280 Upper 270 Middle to

260 Upper Shoreface 250 Cenomanian 240 Bastion Ridge 230 220

210 100.5 Ma 200 190

180

sasel 170

Hassel H 160 Hassel 150

retaceouCs p15 140

Albian Sampled Interval p14 130 Offshore 120 p13 Transition 110

p12 100

90

p11 80 p10 Upper ~113 Ma 70 Offshore

60 Lower Christopher 50 40 p9

r p8 30 p7 Lower 20 p5 p6 Offshore p4 10 p3 Aptian Christophe p1 p2 Formation Palynology Meters Grain size: Clay Silt VF F C samples M Mud sized Sand sized Sandstone Dominated Mudstone Dominated Figure 3: Location of palynological samples and stratigraphy of the Glacier Fiord section modified from Schröder-Adams et al. (2014). Dashed lines represent the sampled interval. Numerical ages for stage boundaries are from ICS (2014).

15 comparison with the Gaudrina irenensis-Trochammina rutherfordi Zone proposed for Arctic

Slope of Alaska (Tappan, 1962; Schröder-Adams et al., 2014). Similarly, dinoflagellate cysts

preserved in the Bastion Ridge Formation indicate a late Albian or Cenomanian age (Núñez-

Betelu et al., 1994; MacRae, 1996).

In addition to the palynology and foraminifera determinations, the Bastion Ridge

Formation is also dated by stratigraphic relationships. For example, at the Strand Fiord locality

on western Axel Heiberg Island, the Bastion Ridge Formation interfingers with the Strand Fiord

Formation, a unit containing basaltic flows (Embry and Osadetz, 1988; Ricketts et al., 1985).

One Ar40/Ar39 radiometric date from upper lava flows of the Strand Fiord Formation at the

Strand Fiord locality give an age of 95.3 +/- 0.2 Ma (Ricketts et al., 1985), which indicates a late

Cenomanian age based on ICS (2014). The Strand Fiord Formation also contains Metasequoia

Miki (Miki 1941) branchlet impressions indicating a Cenomanian age based on the first appearance of this taxon in western Canada, Alaska and Russia (Ricketts et al., 1985; Núñez-

Betelu et al., 1992; Tarduno et al., 1998; LePage et al., 2005). Due to the interfingering

association with the Strand Fiord Formation at the Strand Fiord locality and paleontological

evidence, the Bastion Ridge Formation is taken to be Cenomanian in age (Galloway et al., 2012).

Kanguk Formation

The Kanguk Formation was named in 1963 by ‘Operation Franklin’ geologists for a 364

m succession of marine mudstone on Axel Heiberg Island (Plauchut and Jutard, 1976; Fortier et

al., 1963). It ranges from less than 70 m thick on Ellesmere Island up to about 1000 m thick on

Bylot Island (Miall et al., 1980). The Kanguk Formation conformably overlies the Hassel

16 Formation in the center of the basin and unconformably on the basin margins, such as Ellesmere

Island (Balkwill et al., 1982; Balkwill, 1983; Núñez-Betelu et al., 1994). On Axel Heiberg

Island, where the Bastion Ridge Formation is present between the Hassel and Kanguk

formations, the contacts are both unconformable (Núñez-Betelu et al., 1994).

The Kanguk Formation consists of black mudstone with abundant thin beds of jarositic clay (Balkwill and Hopkins, 1976). The lower beds have oxidation of iron sulphate minerals, and are often extremely acidic (pH 3.5) (Núñez-Betelu, 1994). Frequently a bituminous mudstone unit is present at the base of the lower member (Plauchut and Jutard, 1976; Miall, 1979; Núñez-

Betelu, 1994). Palynomorphs collected from the basal and top beds of the Kanguk Formation indicate that the formation is as old as late Cenomanian, and that the formation ranges to the late

Campanian-early Maastrichtian (Balkwill et al., 1982; Núñez-Betelu, 1994; Hills and Strong,

2007).

METHODS

This study focuses on stratigraphic sections of the Hassel Formation and bounding units at three different localities: Glacier Fiord on southern Axel Heiberg Island and Strathcona and

Cañón fiords on Ellesmere Island. The distance separating these sections is about 400-500 km

(Figure 1). Samples from Glacier Fiord (Figures 3 and 4) were collected as part of a longer stratigraphic section by Jennifer Galloway (GSC-Calgary), Claudia Schröder-Adams (Carleton

University) and Jens Herrle (University of Frankfurt) as part of a GSC organized expedition under the GeoMapping for Energy and Minerals Program – Western Arctic Project (managed by

Keith Dewing, GSC-Calgary). The Glacier Fiord section preserves 1940 m of the Christopher,

Hassel, Bastion Ridge, and Kanguk formations (Schröder-Adams et al., 2014). Samples studied

17 90° 00'

N Stratigraphy

Glacier Glacier AA B Cenozoic Units

78° 30' Kanguk and Bastion Ridge formations

18 Glacier Fiord Hassel Formation

Surprise Fiord Christopher Formation

78° 15'

Other Lower Cretaceous

Km and Jurassic units 0 6 Figure 4: Geological map of the Glacier Fiord area, Axel Heiberg Island showing the sampling location of the Glacier Fiord section (A-B). Map modified from Thorsteinsson and Tozer (1970). Refer to Figure 1 for a regional perspective. � here were collected from the upper Christopher, Hassel, and lower Bastion Ridge formations

(Figure 3). Foraminifera (C. Schröder-Adams) and macrofossil (J. Haggart) biostratigraphy and

carbon isotope stratigraphy of the entire section has already been published (Schröder-Adams et al., 2014; Herrle et al., 2014).

Additional organic rich material from the Hassel Formation was collected from Cañón and Strathcona fiords, Ellesmere Island (Figures 5 and 6) in the early 1990s by Len Hills and

Koldo Núñez–Betelu. Described sections are unavailable, as the location of the field note books is not known. All efforts were made to find the field notebooks but to no avail. Formation information and meterage above the base of section are available and permit placement of the samples and stratigraphic palynological study.

Palynological preparation

Samples were prepared by Linda Dancey at the Geological Survey of Canada, Calgary

Palynological Laboratory following standard extraction techniques (Faegri and Iverson, 1989).

This included acid digestion with hydrochloric acid to remove carbonates and hydrogen fluoride to remove silicates. Heavy liquid separation with zinc bromide was used to separate organic material from mineral matter and oxidation with Schulze’s solution was used to digest non• palynomorph organic matter. Finally, palynomorphs were stained with Safranin O before being mounted with liquid bioplastic (Wood et al. 1996). Palynomorphs were identified using an

Olympus BX61 transmitted light microscope under oil immersion at 400x and 1000x magnification. All microscope slides are stored and curated at the Geological Survey of Canada,

Calgary (Table 2).

19 Table 2: GSC Curation (C-#) for samples in this study Glacier Fiord Depth Strathcona Fiord Depth C-551725 1.5 C-552776 1 C-551726 4 C-552778 6 C-551727 6.5 C-552779 10 C-551728 11 C-552780 13 C-551729 17 C-552781 21 C-551730 19 C-552782 24 C-551731 25 C-552783 27 C-551732 33 C-552784 30 C-551733 36 C-552785 33 C-551734 76 C-552786 36 C-551735 79 C-552788 46 C-551736 99 C-552789 52 C-551737 117.5 C-552790 53 C-551738 131 C-552791 57 C-551739 143.5 C-552792 60 C-551740 286 C-552793 65 C-551741 286.2 C-552794 67 C-551742 286.5 C-552795 70 C-551743 286.6 C-552796 73 C-551744 288 C-552797 76 C-552798 79 C-552799 85 C-552800 94 Cañón Fiord Depth C-552801 120 C-552820 4 C-552802 123 C-552821 9 C-552803 126 C-552822 11 C-552804 129 C-552823 15 C-552805 132 C-552824 20 C-552806 135 C-552826 35 C-552808 141 C-552827 38 C-552809 144 C-552828 41 C-552810 147 C-552829 44 C-552811 150 C-552830 50 C-552814 159

20 83° 30' � 79° 55' Stratigraphy N anCa Fiord óñ Cenozoic Units A B Mt. Bridgman

Kanguk Formation

21 Hassel Formation

Christopher Formation

Km 0 6 Other Lower Cretaceous 79° 45' and Jurassic units

Figure 5: Geological map of the Cañón Fiord area, Ellesmere Island showing the sampling location of the Cañón Fiord section (A-B). Map modified from Thorsteinsson and Tozer (1970). Refer to Figure 1 for a regional perspective. Other Mesozoic and Paleozoic units Hassel Formation Christopher Formation Cenozoic Units Kanguk Formation Stratigraphy

rd Fio 83° 30' aStrathcon 6 Km 0 N

B A 84° 45' 78° 35' 78° 40' Figure 6: Geological map of the Strathcona Fiord area, Ellesmere Island showing the sampling location of the Strathcona Fiord Strathcona Fiord area, Ellesmere Island showing the sampling location Figure 6: Geological map of the a regional perspective. (1970). Refer to Figure 1 for Tozer and Thorsteinsson section (A-B). Map modified from

22 Palynology

A total of 76 samples were examined for palynomorphs. Of this total, 20 samples are from Glacier Fiord (Figure 3), 11 samples are from Cañón Fiord, and 45 samples are from

Strathcona Fiord. A minimum of 300 terrestrial palynomorphs were counted for each sample.

The maximum number counted is 425, the minimum number counted is 300 and the median counted is 329 (Appendix I).

Taxonomic Notes

Two pollen genera accommodate reticulate, tricolpate fossil pollen, Tricolpites Cookson

1947 ex Couper 1953 emend, Potonie 1960 and Retitricolpites van der Hammen 1956 ex Pierce

1961. Due to difficulties in distinguishing these two likely synonymous genera and confusion over which has precedence, reticulate, tricolpate dicotyledonous angiosperm pollen are here referred to as tricolpate undifferentiated (Srivastava 1966; Playford, 1971; Galloway et al.,

2012). Bisaccate pollen identification was not attempted, as their Cretaceous taxonomy remains poorly understood (Traverse, 2007). Monosulcate pollen of Ginkgopsida and Cycadopsida are difficult to differentiate to the species level using light microscopy (Hill, 1990) and are rarely possible even with modern material (Wodehouse, 1933). Nonetheless, the genus Cycadopitys

Wodehouse is used to accommodate monosulcate pollen ranging in size from 39-42 µm and 18•

21 µm wide, following the example of the type specimen Cycadopitys follicularis (Wilson and

Webster, 1946). Monosulcate pollen ranging in size from 33-85 µm long and 20-40 µm wide are assigned to the genus Entylissa Naumova 1939 ex Ishchenko 1952 (Burden and Hills, 1989). All other taxonomic authorities are given in Table 3.

23 Table 3: Taxonomic authorities of taxa documented in this study. Taxa Authority Aequitriradites spp. Delcourt and Sprumont 1955 Appendicisporites spp. Weyland and Krieger 1953 Baculatisporites comaumensis (Cookson) Potonie 1956 Biretisporites potoniaei Delcourt and Sprumont 1955 Camarozonosporites ambigens (Fradkina) Playford 1971 Cicatricosisporites hallei Delcourt and Sprumont 1955 Cicatricosisporites spp. Potoniie and Gelletich 1933 Cicatricososporites spp. Thomson and Pflug 1953 Cingutriletes clavus (Balme) Dettmann 1963 Cingutriletes spp. Pierce 1961 Concavisporites juriensis Balme 1957 Concavissimisporites spp. Delcourt and Sprumont 1955 Converrucosisporites spp. Potonie and Kremp 1954 Cyathidites australis Couper 1953 Cyathidites minor Couper 1953 Cycadopites spp. Wodehouse 1933 Deltoidospora hallii Miner 1935 Deltoidospora psilostoma Rouse 1959 Deltoidospora spp. Miner 1935 Entylissa spp. Samollovich 1963 Equesetosporites type D Steves and Barghorn 1959 Gleicheniidites circinidites (Cookson) Brenner 1963 Gleicheniidites senonicus Ross 1949 Laevigatosporites ovatus Wilson and Webster 1946 Laricoidites magnus (Potonie) Potonie, Thompson, and Thiergart 1950 Lycopodiumsporites expansus Singh 1971 Lycopodiumsporites spp. Thiergart ex Delcourt and Sprumont 1955 Monocolpate undifferentiated N/A Murospora spp. Somers 1952 Neoraistrickia truncata (Cookson) Potonie 1956 Neoraistrickia spp. Potonie 1956 Osmundacidites wellmannii Couper 1953 Pilosisporites trichopapillus Delcourt and Sprumont 1955 Pilosisporites spp. Delcourt and Sprumont 1955 Podocarpites spp. Bolkhovitina 1986 Sciadopityspollenites spp. Raatz ex Potonie 1958 Stereisporites antiquasporites (Wilson and Webster) Dettmann 1963 Stoveresporites spp. Norvick and Burger 1975 Taxodiaceaepollenites hiatus (Potonie) Kremp 1949 Tricolpate undifferentiated N/A Trilobosporites marylandensis Brenner 1963 Undifferentiated bisaccate pollen N/A Undulatisporites spp. Thomson and Pflug 1953 Verrucosisporites rotundus Singh 1964 Verrucosisporites spp. Oybova and Jachowicz 1957 Vitreisporites pallidus (Reissinger) Nilsson 1958

24 Multivariate Statistical Analyses

To delineate which taxa co-occur regionally, palynological data from all sections were

combined for use in R-mode hierarchical cluster analysis. This analysis employed Ward’s

minimum variance method and Euclidian distance on relative abundance data using the computer

program SYSTAT®13. Cluster analysis is a statistical process of grouping sets of variables (in this case taxa) into clusters in such a way that variables in each cluster are more similar to each other than to those in a different cluster. This involves a measurement of distance (in these case co-occurrences between objects (Euclidian or squared Euclidian distances) and a method of grouping these objects (Ward’s method). Ward’s method and Euclidian distance were chosen because these are known for reliably creating distinctive clusters in palynological data (Birks and

Gordon, 1985; Kent and Cooker, 1992; Hills and Strong, 2007; Galloway et al., 2013). Q-mode hierarchical cluster analysis was also applied to explore relationships between samples based on their palynomorph content and to test the hypothesis “Samples from Hassel Formation, irrespective of location (Glacier Fiord, Strancona Fiord and Cañón Fiord) contain similar palynoassemblages.”

Multidimensional scaling ordination was performed using the computer program

SYSTAT®13 as a means to identify populations of taxa based on palynological composition for comparison with R-mode cluster analysis. Many ordination techniques exist (principal components analysis (PCA), correspondence analysis (CA), multidimensional scaling (MDS)) and are used to represent multivariate data by reducing the number of variables to the two or three principal variables that explain the majority of variation in the data. Multidimensional scaling was chosen for this study because MDS does not contain underlying assumptions that

25 PCA or CA methods assume about species distribution patterns along a compositional gradient

(Kruskal, 1964; Prentice, 1977; Minchin, 1987; Clarke, 1993).

Palynoassemblages were plotted stratigraphically using the program Tilia to view

changes over time (Grimm, 1993-2001). A stratigraphically constrained incremental sum of

squares cluster analysis (CONISS) based on square root transformed (to up-weigh rare taxa)

relative abundance data of palynomorphs from obligately terrestrial plants was also added using

Tilia. This was done to aid the delineation of pollen and spore assemblage zones for each of the

three sections and to observe important changes in palynomorph assemblage composition over

time (Grimm, 1987; 2004).

RESULTS

A variety of Early Cretaceous pollen and spores typical of previous Hassel Formation studies are preserved at Glacier, Cañón and Strathcona fiords (Plate 1) (Hopkins and Balkwill,

1973; Galloway et al., 2012). A total palynomorph count of 6342 was achieved (Mean: 317,

Standard Deviation: 17) for Glacier Fiord samples, 11,431 (Mean: 336, Standard Deviation: 30)

for Strathcona Fiord samples, and 3,110 (Mean: 311, Standard Deviation: 20) for Cañón Fiord

samples. Palynomorphs identified in samples from these three stratigraphic sections represent 47

different pollen and spore taxa (Table 3).

Angiosperm pollen

A single monocotyledonous pollen grain was recovered from the upper Christopher

Formation at Glacier Fiord and two were recovered from the Hassel Formation at Strathcona

26 PLATES Photomicrographs of pollen and spores captured using differential interference contrast oil immersion preserved in upper Christopher, Plate 1: Hassel, and Bastion Ridge formations from exposures on Ellesmere Axel Heiberg islands. Photograph scale bars represent 10 μm. See following page for explanation.

27 Explanation of Plate 1: GSC-Calgary Curation number (C-number), GSC Calgary Palynology Laboratory Preparation number (P-number), Sample number (SF = Strathcona Fiord section; CF = Cañón Fiord section; GF = Glacier Fiord section), GSC Specimen number, and England Finder coordinates.

1. Dicot tetrad. C-552818 / P5257-44B / SF-11 / GSC specimen / U12 2. Dicot pollen. C-552823 / P5257-49B / CF-15 / GSC specimen / F14-2 3. Dicot pollen. C-552828 / P5257-41B / CF-41 / GSC specimen / P13 4. Dicot pollen. C-552826 / P5257-52B / CF-35 / GSC specimen / H13-4 5. Dicot pollen. C-551732 / P5256-8B / GF-33 / GSC specimen / K14-4 6. Monocot pollen. C-552819 / P5257-45B / SF-42.5 / GSC specimen / G13-4 7. Monocot pollen. C-552819 / P5257-45B / SF-42.5 / GSC specimen / G15-1 8. Monocot pollen. C-551726 / P5256-2B / GF-4 / GSC specimen / R18-3 9. Baculatisporites comaumensis. C-552805 / P5257-31B / SF-132 / GSC specimen / N19-4 10. Stoverisporites spp. C-552801 / P5257-27B / SF-120 / GSC specimen / K44-3 11. Gleicheniidites cf. G. circinidites. C-552806 / P5257-21B / SF-70 / GSC specimen / T12- 4 12. Gleicheniidites senonicus. C-552801 / P5257-27B / SF-120/ GSC specimen / P26-3 13. Cicatricosisporites spp.C-552778 / P5257-4B / SF-6 / GSC specimen / U18-4 14. Biretisporites potoniaei. C-552801 / P5257-27B / SF-120 / GSC specimen / O41 15. Cyathidites spp. C-552780 / P5257-6B / SF-13 / GSC specimen / T42-3 16. Murospora spp. C-552776 / P5257-2B / SF-1 / GSC specimen / H17 17. Pilosisporites trichopapillus. C-552776 / P5257-2B / SF-1 / GSC specimen / W11 18. Converrucosisporites spp. C-552801 / P5257-27B / SF-120 / GSC specimen / Q25-2 19. Trilobosporites marylandensis. C-552811 / P5257-37B / SF-150 / GSC specimen / M13- 4 20. Concavissimisporites spp. C-552776 / P5257-2B / SF-1 / GSC specimen / S23 21. Appendicisporites spp. C-552782 / P5257-8B / SF-24 / GSC specimen / F16 22. Neoraistrickia truncata. C-552776 / P5257-2B/ SF-1 / GSC specimen / H14 23. Lycopodiumsporites expansus. C-552820 / P5257-46B / CF-1 / GSC specimen / J11-4 24. Camarozonosporites ambigens. C-552818 / P-5257-44B / SF-11 / GSC specimen / 25. Laevigatosporites ovatus. C-552801/ P5257-27B / SF-120 / GSC specimen / T30-1 26. Aequitriradites spp. C-552780 / P5257-6B / SF-13 / GSC specimen / T10-2 27. Cingutriletes clavus. C-551741 / P5256-17B / GF-286.2 / GSC specimen / P17 28. Stereisporites antiquasporites. C-552801 / P5257-27B / SF-120 / GSC specimen / O15-1 29. Sciadopityspollenites spp. C-552801 / P5257-27B / SF-120 / GSC specimen / Q14-1 30. Equesetosporites type D. C-552793 /P5257-19B / SF-64.5 / GSC specimen / V11-1 31. Entylissa spp. C-552801/ P5257-27B / SF-120 / GSC specimen / V21-3 32. Cycadopitys spp. C-552801/ P5257-27B / SF-120 / GSC specimen / U16 33. Taxodiaceaepollenites hiatus (small grain) / Laricoidites magnus (larger grain) C- 552814/ P5257-40B / SF-159 / GSC specimen / H23-3 34. Undiff. Bisaccate pollen, C-552788 / P5257-14B / SF-46/ GSC specimen / G18-2 35. Podocarpites spp. C-552780 / P5257-6B / SF-13 / GSC specimen / W10-3 36. Vitreisporites spp. C-552801 / P5257-27B / SF-120 / GSC specimen / W20-2

28 Fiord (Plate 1). A total of 8 reticulate, tricolpate angiosperm pollen were also recovered: one from the MacDougall Point Member of the Christopher Formation at Glacier Fiord, three from the Hassel Formation at Cañón Fiord and four, in the form of a single tetrad, from Strathcona

Fiord.

R-Mode Cluster Analysis

Hierarchical cluster analysis of all sections reveals at least five informal palynoassemblages (Figure 7). Each Assemblage is named after the most dominant plant groups.

Pteridophytina-Podocarpaceae assemblage

This assemblage is dominated by Pteridophytina spores (Deltoidospora hallii, mean

0.797 +/- 1.053 SD %; Cicatricosisporites spp., mean 0.043 +/- 0.144; Murospora spp., mean

0.029 +/- 0.121 SD %; Stoverisporites spp., mean 0.005 +/- 0.038 SD %; SD % of each sample) that collectively represent 94% of Pteridophytina-Podocarpaceae assemblage. Minor amounts of

Podocarpites spp. (mean 0.052 +/- 0.148 SD %), and monocotyledonous angiosperm (mean

0.005 +/- 0.038 SD %) pollen each represent 5.5% and 0.5% of the Pteridophytina-

Podocarpaceae assemblage, respectively. The Pteridophytina-Podocarpaceae assemblage represents only one percent of the total palynological population of all samples from all sections.

Pteridophytina-Bryophyta assemblage

Pteridophytina spores (Gleicheniidites senonicus, mean 7.378 +/- 4.305 SD %;

Biretisporites potoniaei, mean 2.683 +/- 1.765 SD %; Gleicheniidites spp., mean 2.655 +/- 1.807

SD %; Cicatricosisporites spp., mean 0.497 +/- 0.689 SD %; Baculatisporites comaumensis,

29 5 4 3 Euclidian Distances Cluster Tree Tree Cluster 2 1 0 spp. spp. spp. spp. spp. spp. spp. spp. spp. spp. spp. spp. spp. spp. spp. spp. type D Entylissa spp. Murospora spp. Tricolpate undiff. Tricolpate Cycadopites Cingutriletes Cyathidites minor spp. Vitreisporites Podocarpites Pilosisporites Neoraistrickia Deltoidospora Aequitriradites Deltoidospora hallii Deltoidospora Gleicheniidites Monocolpate undiff. Cingutriletes clavus Laricoidites magnus Cyathidites australis Stoveresporites spp. Undulatisporites Taxa Taxa spp. Verrucosisporites Neoraistrickia truncata Appendicisporites Biretisporites potoniaei Cicatricosisporites Undiff. bisaccate pollen Undiff. Cicatricososporites Cicatricosisporites hallei Equesetosporites Gleichinidites circinidites Lycopodiumsporites Concavisporites juriensis Laevigatosporites ovatus Deltoidospora psilostoma Gleicheniidites senonicus Sciadopityspollenites Converrucosisporites Concavissimisporites Osmundacidites wellmanii Verrucosisporites rotundus Verrucosisporites Pilosisporites trichopapillus Taxodiaceaepollenites hiatus Taxodiaceaepollenites Stereisporites antiquasporites Trilobosporites marylandensis Trilobosporites Lycopodiumsporites expansus Lycopodiumsporites Baculatisporites comaumensis Camarozonosporites ambigens Taxaceae Taxaceae Bisaccate Laricoidites Bryophyta Assemblage Assemblage Cupressaceae Pteridophytina Pteridophytina Pteridophytina Podocarpaceae Sciadopityspollenites Bryophyta 3% Pteridophytina 2% 9% Lycopodiopsida Lycopodiopsida or 94% 83% CT 6% 81% 18% 95% 84% 13% 7% Laricoidites Bryophyte Bisaccate 3% Laricoidites Pteridophytina 2% Pteridophytina Pteridophytina Podocarpaceae Bisaccate Sciadopityspollenites Taxaceae Taxaceae Bryophyta Podocarpace Pteridophytina Sciadopityspollenites Pteridophytina Cupressaceae Composition Composition Taxa with <1%: Equisetaceae Taxa Cycadopsida Taxa with <1%: Monocolpate pollen Taxa Taxa with <1%: Lycopodiopsida with <1%: Lycopodiopsida Taxa Ginkgopsida Lycopodiopsida Lycopodiopsida Taxa with <1%: Pteridosperm, Lycopodiopsida with <1%: Pteridosperm, Lycopodiopsida Taxa Taxa with <1%: Pteridophytina, Tricolpate pollen with <1%: Pteridophytina, Tricolpate Taxa Figure 7: R-mode hierarchical cluster analysis of all taxa showing the five resultant assemblages. Pie charts to the left analysis of all taxa showing the five resultant assemblages. Figure 7: R-mode hierarchical cluster plant groupings, as discussed in the text. Plotted show the composition of these assemblages in terms of their major using Systat®13.

30 mean 0.406 +/- 0.672 SD %; Camarozonosporites spp., mean 0.059 +/- 0.214 SD %;

Appendicisporites spp., mean 0.014 +/- 0.062 SD %;) collectively represent 81% of

Pteridophytina-Bryophyta assemblage. Bryophyta spores (Steresisporites antiquasporites, mean

3.166 +/- 5.031 SD %; Cinguitriletes spp., mean 0.005 +/- 0.041 SD %) represent 18.5% of this assemblage, while Pteridospermatophyta pollen (Vitreisporites spp., mean 0.083 +/- 0.280 SD

%) and Lycopodiopsida spores (Lycopodiumsporites expansus, mean 0.025 +/- 0.116 SD %) make up the remaining 0.5%. Pteridophytina-Bryophyta assemblage represents 17% of the total palynological population.

Cupressaceae-Taxaceae assemblage

Taxodiaceaepollenites hiatus (representing families Cupressaceae and Taxaceae) pollen predominates (mean 46.515 +/- 18.404 SD %), representing 95% of this assemblage. Pollen with

Cycadopsida/Ginkgopsida affinities (Cycadopites spp., mean 1.336 +/- 1.495 SD %) and

Lycopodiopsida spores (Lycopodiumsporites spp., mean 0.938 +/- 1.291 SD %) make up 2.8% and 2% of the assemblage, respectively. Pteridophytina spores (Gleicheniidites circinidites, mean 0.035 +/- 0.134 SD %; Concavisporites juriensis, mean 0.031 +/- 0.128 SD %;

Concavissimisporites spp., mean 0.015 +/- 0.095 SD %;) and dicotyledonous angiosperm pollen

(mean 0.031 +/- 0.127 SD %) make up the remaining less than one percent. Cupressaceae-

Taxaceae assemblage represents 49% of the total palynological assemblage.

Bisaccate assemblage

This assemblage is characterized by 84% undiff. bisaccate pollen (mean 26.588 +/•

17.533 SD %), 11% Pteridophytina spores (Cyathidites minor, mean 0.940 +/- 1.030 SD %;

31 Osmundacidites wellmanii, mean 0.672 +/- 0.978 SD %; Verrucosisporites spp., 0.474 mean +/•

0.733 SD %; Cyathidites australis, mean 0.442 +/- 0.483 SD %; Laevigatosporites spp., 0.370 mean +/- 0.505 SD %; Deltoidospora spp., 0.211 mean +/- 0.571 SD %; Pilosisporites spp.,

0.116 mean +/- 0.264 SD %; Converrucosisporites spp., mean 0.107 +/- 0.235 SD %;

Cicatricosisporites halleii, mean 0.105 +/- 0.249 SD %; Verrucosisporites rotundus, mean 0.040

+/- 0.185 SD %; Undulatisporites spp., mean 0.025 +/- 0.165 SD %; Trilobosporites marylandensis, mean 0.009 +/- 0.052 SD %), 3% Bryophyta spores (Cingutriletes clavus, mean

0.990 +/- 1.894 SD %; Aequitriradites spp., mean 0.029 +/- 0.107 SD %), 1.5% pollen with

Gingkopsida and/or Cycadopsida affinities (Entylissa spp., mean 0.452 +/- 0.728 SD %) and less than 1% Lycopodiopsida spores (Neoraistrikia spp., mean 0.023 +/- 0.098 SD %). This assemblage represents 32% of the total palynological population.

Sciadopityspollenites-Laricoidites assemblage

This assemblage consists of 83% conifer pollen (Sciadopityspollenites spp., mean 1.032

+/- 1.568 SD %; Laricoidites magnus, 0.299 mean +/- 0.597 SD %), this assemblage also contains 9% Pteridophytina spores (Deltoidospora psilostoma, mean 0.113 +/- 0.250 SD %;

Pilosisporites trichopapillus, mean 0.029 +/- 0.105 SD %), 7.5% Lycopodiopsida spores

(Neoraistrikia truncata, mean 0.124 +/- 0.209 SD %), and less than 1% spores with affinities to horsetails (Equesetosporites type D, mean 0.009 +/- 0.051 SD %). This assemblage represents

1% of the total palynological population.

32 Multidimensional Scaling Ordination

Multidimensional Scaling Ordination presented in Figure 8 independently reveals a similar clustering of taxa to that found in the R-Mode Cluster Analysis presented in Figure 7. I interpret the first dimension (x-axis) to represent moisture content and the second dimension (y- axis) to represent disturbance (see Discussion section). The first axis shows a spread of lowland, moisture-loving Pteridophytina-Podocarpaceae and Pteridophytina-Bryophyta assemblages to the left, with drier, upland Bisaccate, and Sciatopityspollenites-Laricoidites assemblages to the right.

The second axis places Pteridophytina and Bryophyta spores at the top, with tree taxa such as

Laricoidites magnus and Sciadopityspollenites spp. in the middle and Taxodiaceaepollenites hiatus near the bottom.

CONISS

The relative abundance of pollen and spores are grouped into R-mode clusters and plant groups, and both are plotted stratigraphically to view trends over time at each of the three sections (Figures 9 to 11). Informal stratigraphic zones are delineated using CONISS and discussed below for each of the three sections. CONISS was run on pollen and spore relative abundances, while R-mode clusters were included in Figures 9 to 11 for comparison purposes only.

Glacier Fiord

At Glacier Fiord, five stratigraphically constrained sample clusters are identified, labelled

GF-A though GF-E (Figure 10). By comparing these clusters, ecological information may be obtained. GF-A, the lowermost cluster that includes samples from the upper Christopher and

33 R-Mode Assemblages Pteridophytina-Podocarpaceae Pteridophytina-Bryophyta Cupressaceae-Taxaceae Bisaccate Deltoidospora spp. Sciadopityspollenites-Laricoidites Verrucosisporites spp. Lower Disturbance 1 Vitreisporites spp.

Neoraistrickia truncata Cyathidites australis

Gleichinidites senonicus Dictyophylladites harissii Osmundacidites wellmanii

Cicatricosisporites spp. Cingutriletes spp. Laevigatosporites ovatus Lycopodiumsporites expansus Baculatisporites comaumensis Cyathidites minor

Verrucosisporites rotundus Cingutriletes clavus Biretisporites potoniaei Undifferentiated bisaccate pollen

Neoraistrickia spp. Entylissa spp.

Appendicisporites spp. Converrucosisporites spp. Undulatisporites spp. Camarozonosporites ambigens

0 Stereisporites antiquasporites Aequitriradites spp. Deltoidospora psilostoma

Equesetosporites type D Laricoidites magnus Sciadopityspollenites spp. Podocarpites spp.

Pilosisporites trichopapillus Concavisporites juriensis Stoveresporites spp. Monocolpate pollen High Disturbance Murospora spp. Cicatricosisporites hallei

Trilobosporites marylandensis

Cicatricososporites spp. Cycadopites spp. Deltoidospora hallii Concavissimisporites spp. Gleichinidites circinidites

Lycopodiumsporites spp.

-1 Taxodiaceaepollenites hiatus Pilosisporites spp.

Tricolpate pollen High moisture Low moisture -1 0 1

Figure 8: Multidimensional Scaling Ordination (MDS). Note that both axes are unitless. Assemblages defined using R-mode cluster analysis (Figure 7). Plotted using Systat®13. �

34 R-Mode Assemblages

FormationHeight abovePteridophytina-Podocarpaceae basePteridophytina-Bryophyta (m) Cupressaceae-Taxaceae Bisaccate Sciadopityspollenites-LaricoiditesCONISS Zones CONISS B.R. 290 GF-E 280 270 260 250 240

230 a a

220 at

210 D o o

200 N 190 180 170 160 150 140 130 120 GF-D Hassel Formation Formation Hassel 110 100 90 GF-C 80

70 ta ta 60

50 o Da o

40 N 30 GF-B 20 Ch. 10 GF-A 0 20 40 20 40 60 20 10 " 0.0 0.2 0.4 0.6 0.8 1.0 1.2 Total sum of squares Relative abundance (%) Figure 9: Stratigraphically constrained incremental sum of squares cluster analysis (CONISS) of Glacier Fiord samples showing stratigraphic zones GF-A through GF-E. Plotted using Tillia (Grimm, 2001).

35 R-mode Assemblages

FormationHeight above base (m) CONISS Pteridophytina-PodocarpaceaePteridophytina-Bryophyta Cupressaceae-Taxaceae Bisaccate Sciadopityspollenites-LaricoiditesCONISS Zones 160

150 SF-1 140

130

120

110

100 SF-2

90

80

70

Hassel Formation 60 SF-3

50

40

30 SF-4

20

10 SF-5 0 10 20 20 40 60 20 40 60 80 20 0 1 2 3 4 Relative abundance (%) Total sum of squares Figure 10: Stratigraphically constrained incremental sum of squares cluster analysis (CONISS) of Strathcona Fiord samples showing stratigraphic zones SF-A through SF-E. Plotted using Tillia (Grimm, 2001).

36 R-mode assemblages

CONISS Zones FormationHeight Pteridophytina-Podocarpaceae above basePteridophytina-Bryophyta (m) Cupressaceae-Taxaceae Bisaccate Sciadopityspollenites-Laricoidites CONISS 50

45 CF-C

40

35

30 ta

a

D o o

25 N Hassel Formation Formation Hassel

20

15 CF-B

10 CF-A

5

10 20 40 60 20 40 60 80 20 0.0 0.2. 0.4 0.6 0.8 1.0 Relative abundance (%); shaded areas represent 5% exaggeration of selected taxa. Total sum of squares Figure 11: Stratigraphically constrained incremental sum of squares cluster analysis (CONISS) of Cañón Fiord samples showing stratigraphic zones CF-A through CF-C. Plotted using Tillia (Grimm, 2001).

37 lowermost Hassel formations is characterized by an increasing trend in the relative abundance of

Cupressaceae-Taxaceae assemblage (from 40 to 60%), while the Bisaccate assemblage falls from

25% to 15%. GF-B and GF-C includes samples from the Hassel Formation, are characterized by stabilization in the relative abundance of taxa groups. Zones GF-B and GF-C cluster separately because data does not exist between them. GF-D, also containing samples of Hassel Formation, is characterized by an increase in the relative abundance of Cupressaceae-Taxaceae assemblage

(from 45% to 55%), and a decrease in Pteridophytina-Bryophyta assemblage (from 35% to

25%). GF-E, which includes samples from the Bastion Ridge Formation, is characterized by relatively high abundances of Cupressaceae-Taxaceae Assemblage (up to 65%), Bisaccate assemblage (up to 25%) and Sciadopityspollenites-Laricoidites assemblage (up to 10%).

Strathcona Fiord

At Strathcona Fiord (Figure 11), five informal stratigraphic CONISS palynological zones labelled SF-A through SF-E are delineated using CONISS and visual inspection. Beginning at the base, SF-A is characterized by relatively high percentages of Bisaccate assemblage (60•

80%), relatively low percentages of Cupressaceae-Taxaceae assemblage (10-20%) and

Sciadopityspollenites-Laricoidites assemblage (<5%). SF-B is similar to SF-A, but with relatively higher representation of the Cupressaceae-Taxaceae assemblage (up to 40%), and relatively less Bisaccate assemblage (down to 45%). SF-C is characterized primarily by an increase in Sciadopityspollenites-Laricoidites assemblage (up to 10%) and Pteridophytina-

Podocarpaceae assemblage (up to about 3%), with a slight decrease in Bisaccate assemblage

(down to 20-30%). SF-D is characterized by a large increase in Cupressaceae-Taxaceae assemblage (up to 60%), and a decrease in Bisaccate assemblage (down to 30-40%) and

38 Pteridophytina-Podocarpaceae Assemblage (down to <1%). SF-E is a stabilization of the SF-D flora.

Cañón Fiord

At Cañón Fiord (Figure 12), initial conditions are characterized by decreasing representation of the Pteridophytina-Bryophyta assemblage (from 60 to 30%) and an increasing trend in Cupressaceae-Taxaceae assemblage (30-60%). Pteridophytina and Bryophyta spores and

Cycadopsida pollen decline in relative abundance in this zone, while Cupressaceae-Taxaceae and conifer (Bisaccate undiff, Sciadopityspollenites-Laricoidites) pollen increase. Zones CF-B and

CF-C are characterized by stabilization in the relative abundance of taxa clusters, but with an

increase in Sciadopityspollenites-Laricoidites assemblage (up to 10%). CF-B and CF-C cluster as

separate due to missing data between the two zones.

Q-mode Cluster Analysis

Cluster analysis of all samples from all sections (Q-mode) was used to test the hypothesis

“Samples from Hassel Formation, irrespective of location contain similar palynoassemblages.”

Three informal sample assemblages were revealed and named after sections to which the

predominant representations of samples were collected (Figure 12).

The Glacier Fiord assemblage contains all samples from Glacier Fiord, except one

(sample G286.5 from the Bastion Ridge Formation that maps to the Strathcona Fiord

assemblage), irrespective of formation: Christopher Formation (samples G1.5 through G33),

Hassel Formation (G36 through G143.5) or Bastion Ridge Formation (G286 through G288). The

39 4 3 Cluster Tree Tree Cluster Euclidian Distances 2 1 0 S1 S6 C4 C9 G4 C11 C11 S21 G11 G11 C41 S52 S24 S30 S94 S10 S60 S70 S13 S76 S33 S46 S36 S27 S73 S67 S57 S79 S85 C44 C50 C38 C29 C35 C15 G76 G17 G36 G33 G19 G79 G99 G25 G1.5 G6.5 S141 G131 S132 S144 S120 S150 S123 S126 S147 S129 S159 S135 G286 G288 S64.5 S52.5 G117.5 G117.5 G286.2 G286.6 G286.5 G143.5 Sample + Fiord Fiord fiords Cañón Glacier Strathcona Strathcona Assemblage Pteridophytina-Bryophyta Other 66% 52% 34% 13% 19% 49% 22% R-Mode Assemblages R-Mode 28% Composition Composition 10% Cupressaceae-Taxaceae Cupressaceae-Taxaceae Bisaccate Figure 12: Q-mode hierarchical cluster analysis showing samples from Glacier Fiord and lower Strathcona analysis showing samples from Glacier Fiord and lower Strathcona Figure 12: Q-mode hierarchical cluster their palynological content. Cañón and upper Strathcona Fiord cluster Fiord cluster separately based on in their palynological content. R-mode assemblages present for comparison. suggesting similarities together, Plotted using Systat®13.

40 Glacier Fiord assemblage also includes the lowermost sample from Cañón Fiord (C4), and the second to last sample from Strathcona Fiord (S147). The Glacier Fiord sample assemblage is composed primarily of Cupressaceae-Taxaceae assemblage (52%), Pteridophytina-Bryophyta assemblage (28%) and Bisaccate assemblage (19%).

Cañón Fiord sample assemblage clusters all samples from Cañón Fiord (C9 through

C50), except the lowermost sample (C4) that maps to the Glacier Fiord sample assemblage. The

Cañón Fiord sample assemblage also includes samples from Strathcona Fiord (S73, S123 through S144, and S159). Cañón Fiord sample assemblage is composed primarily of

Cupressaceae-Taxaceae assemblage (66%), Bisaccate assemblage (22%), and Pteridophytina-

Bryophyta assemblage (10%).

Strathcona Fiord assemblage clusters samples S(Strathcona Fiord)1 to S70, S76 to S120,

S147, and G(Glacier Fiord)285 to 286. The Strathcona Fiord assemblage is composed primarily of Bisaccate assemblage (49%), Cupressaceae-Taxaceae assemblage (34%), and Pteridophytina-

Bryophyta assemblage (13%).

Because samples generally plot with other samples from the same section, I reject the null hypothesis that “samples from Hassel Formation, irrespective of location (Glacier Fiord,

Strancona Fiord and Cañón Fiord) contain similar palynoassemblages”. Instead, each section contains a distinct flora.

41 DISCUSSION

Stratigraphic Palynology

When making internationally meaningful age interpretations for formations in the

Sverdrup Basin, one must compare these ages with internationally defined Global Boundary

Stratotype Section Points (GSSP). Unfortunately, at the present time, Lower Cretaceous GSSP’s have not been formally defined. In the case of the Upper Cretaceous, most stages have been defined (ICS, 2014). Of particular interest to this study, the base of the Cenomanian (base of the

Upper Cretaceous) has been located 36 m below the top of the Marnes Bleues Formation on the south side of Mont Risou, Haute-Alpes, France, coinciding with the first appearance of the planktonic foraminifer Rotalipora globotruncanoides Sigal, 1948 (Kennedy et al., 2004).

Furthermore, the base of the Cenomanian is taken to be 100.5 Ma (ICS, 2014). While there is

still some debate on the definition of the Albian-Cenomanian (Lower and Upper Cretaceous)

GSSP, for the purposes of this study the current GSSP as proposed by the ICS will be followed

(Scott et al., 2009; Scott, 2014).

Previous to the ICS formal definition of the Albian-Cenomanian boundary GSSP in 2004,

angiosperm pollen were used to biostratigraphically demarcate the uppermost Lower and

lowermost Upper Cretaceous stages of the Western Interior of Canada. In general, angiosperm

pollen of this interval follows four developmental stages in North America (Singh, 1975 p.379).

First, reticulate monocolpate grains appear during the Aptian in the eastern U.S. and in the early

and middle Albian of the Western Interior of North America. Second, dicotyledonous, reticulate,

tricolpate grains appear in the middle Albian throughout North America. Third, the Albian-

Cenomanian boundary is marked by the appearance of smooth, triangular, tricolporate pollen and

42 permanent tetrads. Fourth, larger tricolporate species with reticulate exines make their entrance

in the middle Cenomanian.

In particular, the “Fish Scale Marker Bed” (FSMB) of the Shaftesbury Formation (Peace

River, northern Alberta), a sandstone or sandstone and siltstone bed containing abundant fish

remains, has frequently been used as the Albian-Cenomanian boundary in the Western Interior

and in subsurface correlations (Singh, 1971; Singh, 1975). The dinoflagellate cyst of latest

Albian age, Ovoidinium verrucosum, occurs between the base of the Shaftesbury Formation and the base of the FSMB (Singh, 1971). Lower Cenomanian palynofloras are recorded above the

FSMB (Singh, 1983). The first appearance of the smooth, triangular, tricolporate angiosperm pollen Nyssapollenites albertensis marks the base of the FSMB, and thus the Albian-

Cenomanian boundary in the Peace River area (Singh, 1971; 1975). This combination of the recently defined international GSSP and the palynologically-based Western Interior regional definitions provide the necessary context for stratigraphic work presented in this study.

Angiosperms in Context

To put Arctic observations into a northern hemispheric context, the first monocolpate angiosperm pollen were found in the Barremian of low to mid-latitudes (0-40°) of the Northern

Hemisphere in the Potomac Group of Maryland, USA, and Wealden Group of southern England

(Couper, 1958; Brenner, 1963; Kemp, 1968; Doyle, 1969; Wolfe et al., 1975; Doyle and Hickey,

1976; Hickey and Doyle, 1977). Middle Albian monocolpate pollen were documented from a number of localities the Swan River Group of Saskatchewan and Manitoba, the Youngstown area and Loon River Formation of southern Alberta, and the Christopher Formation of the Canadian

43 Arctic (Eglinton Island) in the late Albian (Playford, 1971; Singh, 1971; Jarzen and Norris, 1975;

Davies and Wall in Harrison and Brent, 2005).

The first unequivocal tricolpate pollen grains appear around the Barremian-Aptian boundary of multiple Southern Hemisphere sites that includes the Zeweira Formation of southern

Israel (Muller, 1966; Brenner, 1974; Brenner, 1976; Hickey and Doyle, 1977; Traverse, 2007).

The first North American dicotyledonous angiosperms are recorded in the early Albian Potomac

Group of Maryland, USA and subsequently in middle Albian strata of south-eastern Alberta

(Brenner, 1963; Norris, 1967; Doyle, 1969; Playford, 1971; Singh, 1971; Brideaux and

McIntyre, 1975; Wolfe et al., 1975; Doyle and Hickey, 1976). Initially diversity was low with 3 dicotyledonous genera in the middle Albian Potomac Group (Brenner, 1963). Diversity began to increase in the late Albian and Cenomanian up to 8 genera in the Fort St. John Group and 10 genera in the Blairmore Group of southwestern Alberta occur (Singh, 1971; Leckie and Burden,

2001). 16 genera are recorded to the south in the Cheyenne and Kiowa formations of Kansas,

USA (Ward, 1986). The Balyktakh Formation of Arctic Russia document 11 macrofossil genera

(Herman and Spicer, 2010) and 80-85 different angiosperm leaf morphotypes (Herman, 2002).

Cretaceous Alaskan and northeast Asian paleofloras demonstrates that angiosperms were important and perhaps even locally dominant at high latitudes (Smiley, 1969; Spicer and Parrish,

1986; Parrish et al., 1998; Spicer and Herman, 2001; Herman, 2002; Herman and Spicer, 2010).

However, while dicotyledonous angiosperm pollen are recorded in the Sverdrup Basin during the late Albian, these dicotyledonous pollen continue to be rare with <5 genera known in the

44 Cenomanian as shown in this study, Hopkins and Balkwill (1973), Balkwill and Hopkins (1976)

and Galloway et al. (2012).

Sea level was on the rise during the Albian, reaching its highest level during the latest

Cenomanian. This inundation of the continent resulted in the establishment of the Western

Interior Seaway as a southward extension of the Arctic Ocean, which eventually connected to the northward extension of the Gulf of Mexico (Figure 13). As a result, it is likely that the Sverdrup

Basin was relatively isolated from these western and southern areas, likely making comparisons to these regions less meaningful due to potential geographic and genetic separation between early angiosperms. Regional differences associated with continentality may have also played a role in the development of angiosperms in either region (Galloway et al., 2012). For instance, the

Canadian Arctic was found to have a 10°C or lower mean annual temperature based on fossil

wood during the Aptian and Albian (Harland, 2007). On the other hand, Alaska and northeast

Asia were near 13ºC (based on macrofossils) during the latest Albian-Cenomanian due to

moderating effects of the nearby Pacific Ocean (Spicer and Herman, 2010).

Angiosperm Biostratigraphy

A single monocotyledonous pollen grain was recovered from the upper Christopher

Formation at Glacier Fiord, and two grains were recovered from the Hassel Formation at

Strathcona Fiord (Plate 1). A total of 8 reticulate, tricolpate angiosperm pollen from

dicotyledonous plants were recovered in this study: one from the upper Christopher Formation at

Glacier Fiord, three from the Hassel Formation at Cañón Fiord, and four in the form of a single

tetrad from Strathcona Fiord. I compare the palynoassemblages preserved in the three Arctic

45 3

Western Interior Seaway 1 Arctic Ocean 2

Figure 13: Paleogeographic map of North America at the beginning of the Cenomanian (100 Ma). Map was obtained from the ODSN website (http://www.odsn.de/). This map shows the extent of the Western Interior Seaway and the isolation of the Sverdrup Basin (1) from the Balyktakh Formation (2) of Arctic Russia (Herman and Spicer, 2010) and the Nunushuk Formation (3) of Alaska (Spicer and Parrish, 1986) at the beginning of the Cenomanian.

46 sections to the closest well-described Canadian material in the Peace River area of northern

Alberta, spanning the Loon River Formation (middle Albian) through the Dunvegan Formation

(Cenomanian) of Singh (1975).

Using the late Early and early Late Cretaceous regional stage-defining angiosperm

pollen-type stages of Singh (1975), only the first two stages are present in the Arctic material.

Stage 1 is represented by a single monocotyledonous pollen grain of the Christopher Formation,

correlating with an early and middle Albian age in northern Alberta. This appears to correlate

well between Glacier Fiord and the North American Western Interior, indicating that

monocotyledonous plants had also colonized the Canadian High Arctic by at least the early

Albian, but not necessarily in high abundances. This is further supported by reports of

monocotyledonous pollen from the Christopher, Bastion Ridge and Strand Fiord formations

(Núñez-Betelu et al., 1994). However, they are much more common (e.g. 40 figured specimens in Singh, 1971) in more southern locations at this time (Brenner, 1963; Norris, 1967; Singh,

1971, 1983; Hopkins, 1974; Ward, 1986).

The single reticulate tricolpate dicotyledonous angiosperm pollen grains recovered from the upper Christopher Formation at Glacier Fiord and the 7 reticulate tricolpate dicotyledonous angiosperm pollen grains recovered from the late Albian Hassel Formation at Strathcona and

Cañón fiords represent stage 2 of Singh (1975). These pollen grains appear in the middle Albian throughout the Western Interior, a relationship that suggests that there is about a 4 Ma lag time in first occurrences between the Western Interior (middle Albian) and the Canadian High Arctic

(late Albian) localities.

47 Based on the small size, reticulate ornamentation and low abundance of the pollen from the Glacier, Strathcona and Cañón fiords sections, it is likely that early angiosperms of the

Sverdrup Basin were pollinated by insects (Faegri and Iversen, 1989; Traverse, 2007). In general, pollen grains that are dispersed by wind tend to be larger (20-40 μm), lack surface ornament and are present in large quantities (Whitehead, 1969; Muller, 1979; Traverse, 2007). In contrast, those plants pollinated by small insects tend to have smaller pollen (10-20 μm), have simple reticulate or perforate surface sculpture, and produce fewer pollen grains (Ferguson and

Skvarla, 1982; Faegri and Iversen, 1989; Hesse, 2000; Traverse, 2007). All 11 angiosperm pollen

(both dicotyledonous and monocotyledonous) recorded in this study exhibit these characters, and thus are interpreted to have been adapted for insect pollination. Since these pollen types are not adapted for anemophilous long-distance transport, it is possible that the insect pollination strategy of early angiosperms resulted in a relatively slow dispersal time between northern

Alberta and the Sverdrup Basin.

Interestingly, angiosperm pollen have not been recorded from the Bastion Ridge

Formation, which was deposited during the Cenomanian, a period of rapid angiosperm diversification to the south in Alberta (Singh, 1975; 1983; MacRae, 1996; Leckie and Burden,

2001). The Bastion Ridge Formation is time equivalent to the Cenomanian portion of the Hassel

Formation on Ellef Ringnes Island, as described by Galloway et al. (2012). Not only does this suggest a lag time in migration of angiosperms into the Sverdrup Basin, but also a restriction of diversification with respect to more southern flora. However, the Bastion Ridge Formation is a

48 marine mudstone, and pollen may have been subject to sorting effects, potentially reducing or removing the angiosperm pollen signal from the rock record.

It is clear that angiosperms were somehow restricted in their migration into the Canadian

Arctic relative to more southern and western high latitude sites. This may have been due to geographic isolation by the Western Interior Seaway barrier to the west and coupled with cooler temperatures (~10°C or lower mean annual temperature) relative to other Arctic provinces such as Alaska and northeast Asia, which had a mean annual temperature of ~13ºC (Harland, 2007;

Spicer and Herman, 2010). Angiosperms reached the Sverdrup Basin by the late Albian, but were not very common on the landscape at this time, even in lowland environments that would otherwise have been suitable for colonization.

Paleoecology and Paleoclimate

Fossil pollen and spores provide the best evidence of past climate shifts because land plants are sensitive to temperature and moisture changes and often produce copious amounts of chemically resistant pollen and spores (Traverse, 2007). Following the methodology of

Quaternary palynology, paleoclimate parameters can be deduced based on the relative abundances of pollen and spores that have been recovered from Hassel Formation sediments in this study (Faegri and Iversen, 1989). Uncertainty exists regarding ancient ecological tolerances of extinct parent plants in comparison to modern parent plants. However, it is assumed that

Cretaceous paleoenvironments supported taxa with broadly similar ecological tolerances to plants today (Abbink et al., 2004).

49 In general, terrestrial palynomorphs identified from the upper Christopher, Hassel and

Bastion Ridge formations in this study are similar to those previously described in Albian-aged strata in the Sverdrup Basin and central Canada (Hopkins and Balkwill, 1973; Singh, 1971;

Núñez-Betelu et al., 1994, Galloway et al., 2012). This study documents pollen and spores of 7

Pteridophytina (ferns), 3 Bryophyta (mosses), 3 Lycopodiopsida (lycopods), 2 Ginkgopsida

(ginkgos) or Cycadopsida (cycads), and 5 Pinopsida (pines) genera and only a handful of angiosperm pollen (Table 4). These higher taxonomic levels form the basis for ecological interpretations and are discussed below.

Pteridophytina (Ferns)

Most extant Pteridophytina prefer to grow in moist, shady environments. Most often they are tropical, but they are also common in temperate forests and marshes (Abbink et al., 2004).

Therefore, Early Cretaceous Pteridophytina are inferred to have grown in moist, warm conditions along lakes, marshes, or rivers and as understory in forests (Abbink et al., 2004; Schrank, 2010).

In particular, members of Osmundaceae (e.g. Baculatisporites) are often closely associated with

swamp margins, whereas Gleicheniaceae (e.g. Gleicheniidites) and Schizaeceae (e.g.

Cicatricosisporites) are associated with opportunistic colonizing of open and disturbed ground

(Crane, 1987; Greenwood and Basinger, 1994; Collinson, 2002; Page, 2002).

Bryophyta (Mosses, hornworts and liverworts)

Bryophyta, in general, prefer humid, tropical, lowland conditions, but some can withstand

long periods of drought (Glime, 2007). Bryophyta found in this study (primarily Steresiesporites

antiquasporites) likely derived from parent plants growing in humid, lowland environments, and

50 Table 4: Further taxonomic information. Taxa Division Class Order Family Cluster Cicatricososporites spp. Pteridophytina Polypodiopsida Schizaeales Schizaeaceae 1A Deltoidospora hallii Pteridophytina 1A Monocolpate undifferentiated Angiospermae Monocotyledoneae 1A Murospora spp. Pteridophytina 1A Podocarpites spp. Gymnospermae Pinopsida Araucariales Podocarpaceae 1A Stoveresporites spp. Pteridophytina 1A Appendicisporites spp. Pteridophytina Polypodiopsida Schizaeales Schizaeaceae 1B Baculatisporites comaumensis Pteridophytina Polypodiopsida Osmundales Osmundaceae 1B Biretisporites potoniaei Pteridophytina Polypodiopsida Osmundales Osmundaceae 1B Camarozonosporites ambigens Pteridophytina Lycopodiopsida Lycopodiales Lycopodiaceae 1B Cicatricosisporites spp. Pteridophytina Polypodiopsida Schizaeales Schizaeaceae 1B Cingutriletes clavus Bryophytina Sphagnopsida Sphagnales Sphagnaceae 1B Gleicheniidites senonicus Pteridophytina Polypodiopsida Gleicheniales Gleicheniaceae 1B Lycopodiumsporites expansus Pteridophytina Lycopodiopsida Lycopodiales Lycopodiaceae 1B Stereisporites antiquasporites Bryophytina Sphagnopsida Sphagnales Sphagnaceae 1B Vitreisporites pallidus Pteridospermatophyta 1B Concavisporites juriensis Pteridophytina 2A Concavissimisporites spp. Pteridophytina 2A Cycadopites spp. Gymnospermae Cycadopsida Cycadales Cycadaceae 2A Gleicheniidites circinidites Pteridophytina Polypodiopsida Gleicheniales Gleicheniaceae 2A Lycopodiumsporites spp. Pteridophytina Lycopodiopsida Lycopodiales Lycopodiaceae 2A Taxodiaceaepollenites hiatus Gymnospermae Pinopsida Cupressales Cupressaceae 2A Tricolpate undifferentiated Angiospermae Dicotyledoneae 2A Aequitriradites spp. Bryophytina 2B Cicatricosisporites hallei Pteridophytina Polypodiopsida Schizaeales Schizaeaceae 2B Cingutriletes spp. Bryophytina Sphagnopsida Sphagnales Sphagnaceae 2B Converrucosisporites spp. Pteridophytina 2B Cyathidites australis Pteridophytina Polypodiopsida Cyathales Cyatheaceae 2B Cyathidites minor Pteridophytina Polypodiopsida Cyathales Cyatheaceae 2B Deltoidospora spp. Pteridophytina 2B Entylissa spp. Gymnospermae Ginkgoopsida Ginkgoales Ginkgoaceae 2B Laevigatosporites ovatus Pteridophytina Polypodiopsida Polypodiales Polypodiaceae? 2B Neoraistrickia spp. Pteridophytina Lycopodiopsida Selaginellales Selaginellaceae 2B Osmundacidites wellmannii Pteridophytina Polypodiopsida Osmundales Osmundaceae 2B Pilosisporites spp. Pteridophytina 2B Trilobosporites marylandensis Pteridophytina 2B Undifferentiated bisaccate pollen Gymnospermae Pinopsida Pinales Pinaceae 2B Undulatisporites spp. Pteridophytina 2B Verrucosisporites rotundus Pteridophytina 2B Verrucosisporites spp. Pteridophytina 2B Deltoidospora psilostoma Pteridophytina 2C Equesetosporites type D Pteridophytina Equisetopsida Equisetales Equisetaceae 2C Laricoidites magnus Gymnospermae Pinopsida Pinales Pinaceae 2C Neoraistrickia truncata Pteridophytina Lycopodiopsida Selaginellales Selaginellaceae 2C Pilosisporites trichopapillus Pteridophytina 2C Sciadopityspollenites spp. Gymnospermae Pinopsida Pinales Pinaceae 2C

51 probably have affinities to the modern genus Sphagnum L. (peat moss) (Andrus, 1986; Abbink et al., 2004).

Lycopodiopsida (Club mosses, fir mosses)

Like Bryophyta, most members of the class Lycopodiopsida live in humid tropical regions today, but can be found in temperate areas, where they grow in moist settings such as lowlands near rivers (Abbink et al., 2004). Triassic Lycopodiopsida appear to have lived near marine shorelines in delta systems (Retallack, 1975), and were also components of communities dominated by Pteridophytina established around coastal or deltaic lowlands and poorly developed flood plains (Dejax et al., 2007; Stukins et al., 2013).

Gingkopsida and Cycadopsida

Monosulcate pollen identified as Cycadopitys and Entylissa are associated with classes

Cycadopsida and Ginkgopsida. Class Gingkopsida, abundant throughout the Mesozoic (Zhou,

2009), has only one modern representative, Ginkgo biloba L. This species exists largely under cultivation in China and Japan. Ginkgo biloba grow best in areas with annual mean temperature of 10-18 °C and annual mean precipitation of 600-1000 mm (He et al., 1997). Similarly, it appears that members of Ginkgopsida were abundant and diverse in mesic, warm temperate to temperate climates, such as were present during the Jurassic and Early Cretaceous (Del Tredici et al., 1992; He et al., 1997; Zhou, 2009). Expansion and contraction of ginkgo distribution, both in abundance and geographically through geological time appears to follow shifts of the mesic temperate climatic zone (Vachrameev, 1987; 1991). A study of Late Cretaceous and Cenozoic ginkgo fossils found that they inhabited streamside and levee environments, suggesting early

52 successional ecology and shade intolerance (Royer et al., 2003). Extant Cycadopsida occur in

tropical and sub-tropic lowland regions free of frost and can withstand droughts (Abbink et al.,

2004). Therefore, parent Cycadopsida or Ginkgopsida plants represented by Cycadopitys and

Entylissa pollen may have had similar ecological tolerances in drier parts of lowland

environments or on well-drained upslope environments (Abbink et al., 2004).

Pinopsida

Class Pinopsida is represented in this study by families Cupressaceae, Taxaceae,

Pinaceae (including undifferentiated bisaccate pollen and Laricoidites magnus) and

Sciadopityacea. Modern Cupressaceae and Taxaceae are widely distributed throughout North

America and are competitive in moist temperate climates, where they may grow in upland

environments or may form communities in lowland marshes (Abbink et al., 2004; Schrank,

2010; Galloway et al., 2013). Taxodiacean conifers are interpreted to have been hygrophilous

plants that thrived in warm to temperate wet lowland environments (Vakhrameev, 1991; Dejax et al., 2007). Modern Cupressaceae are strong components in upland habitats in moist and temperate climates (Gavin et al., 2005; Galloway et al., 2007).

Parent plants of bisaccate pollen were likely upland conifers growing in cool temperatures and relatively dry upland areas, similar to the modern genera Pinus L. (pine) and

Picea Mill. (spruce) (Abbink et al., 2004; Schrank, 2010; Galloway et al., 2013). Parent plants

of Laricoidites magnus are likely similar to the modern genus Larix Mill. (Larch). They exist

today in the Pacific Northwest, and across the Northwest Territories of Canada (for example

Larix laricina (Du Roi) K. Koch (Tamarack); Uchytil, 1991), where they are common in cool,

53 temperate, mixed forest communities (Arno and Habeck, 1972). One extant representative of the

Family Sciadopityaceae, Sciadopitys verticillata (Thunberg) Siebold et Zuccarini 1842, exists

today in Japan, where it is common in mixed conifer and angiosperm forest communities in

association with Pteridophytina, Lycopodiopsida and Bryophyta growing in cool and moist

environments (Abbink et al., 2004).

R-Mode Cluster Analysis

R-mode cluster analysis was used to delineate five clusters of taxa (Figure 7). I interpret

these clusters to represent palynoassemblages, groups of pollen and spores that tend to co-occur.

Pteridophytina-Podocarpaceae and Pteridophytina-Bryophyta assemblages are dominated by

spore-bearing (Pteridophytina, Bryophyta, Lycopodiopsida) taxa, including the orders

Schizaeales, Osmundales, Gleicheniales, and Sphagnales (Table 4). Today, Osmundales and

Sphagnales are commonly associated with swamps and wetlands, whereas Schizaeales and

Gleicheniales are opportunistic colonizers of open and disturbed ground (Crane, 1987;

Greenwood and Basinger, 1994; Collinson, 2002; Page, 2002). The mixture of pollen and spores

assignable to these plant orders therefore is suggestive of provenance from vegetation growing in

a swampy lowland area, likely proximal to a river where frequent disturbance occurred.

The Cupressaceae-Taxaceae assemblage is dominated by Cupressaceae-Taxaceae pollen, with some Cycadopsida or Ginkgopsida pollen representation (Figure 7). This pollen likely originated from plants growing in a humid, lowland environment where Cupressaceae-Taxaceae parent plants may have formed climax communities (Vakhrameev, 1991; Abbink et al., 2004;

Dejax et al., 2007; Schrank, 2010; Galloway et al., 2013). Alternatively, because modern

54 Cupressaceae-Taxaceae parent plants are also dominant components of hinterland vegetation in temperate and moist climates at present and during the Holocene, pollen assignable to this group may represent a signal from upland trees growing in moist settings (Galloway et al., 2010).

Cycadopsida or Ginkgopsida pollen represents drier parts of lowland or a well-drained upslope environment (Vachrameev, 1987; 1991; Del Tredici et al., 1992; He et al., 1997; Royer et al.,

2003; Abbink et al., 2004; Zhou, 2009).

Both the Bisaccate and Sciadopityspollenites-Laricoidites assemblages are largely dominated by Pinaceae pollen. The Bisaccate assemblage is dominated by undifferentiated bisaccate pollen, which may have originated from a variety of conifer parent plants. For example, the modern genera Picea Mill. and Pinus L. produce bisaccate pollen and often live in cool, temperate upland forests (Stukins et al., 2013). The difference between the two assemblages is the composition: Sciadopityspollenites-Laricoidites assemblage contains a more diverse range of conifers, suggesting more mixed forest upland. Mixed communities are often the result of regional fires, and therefore I interpret the Sciadopityspollenites-Laricoidites assemblage to represent evidence of fires in the region (Ahlgren and Ahlgren, 1960; Rowe and

Scotter, 1973).

Dicotyledonous angiosperm pollen from samples of Hassel Formation exposed at Cañón

Fiord cluster within the Cupressaceae-Taxaceae assemblage, which I interpret to be lowland or upland community composed primarily of Cupressaceae-Taxaceae, Cycadopsida/Ginkgopsida and Gleichenales. The Gleichenales are a family that includes members able to grow in open environments (Crane, 1987; Collinson, 2002; Page, 2002). The association with Cupressaceace•

55 Taxaceae and also suggests that early angiosperms in the Canadian Arctic were able to grow adjacent to lowland and potentially moist upslope communities. This provides evidence to support previous interpretations of early angiosperms being competitive in disturbed lowland environments near water, such as in riparian or deltaic environments, or potentially evidence that dicotyledonous angiosperms were inhabiting upland environments (Hickey and Doyle, 1977;

Retallack and Dilcher, 1986; Herman, 2002; Coiffard et al., 2006; Royer et al., 2010). Therefore, this pollen and spore assemblage likely reflects a mosaic of lowland boggy habitats and surrounding open environments, and potentially moist upslope environments.

The single monocot pollen from the Christopher Formation at Glacier Fiord clusters in the Pteridophytina-Podocarpaceae assemblage, which I interpret as a swampy lowland environment composed mostly of Pteridophytina and Podocarpaceae. Therefore this association would indicate that monocots in the Sverdrup Basin were adapted to life in moist, open habitats near, or on the fringes of a swamp or marshland environment.

Multidimensional Scaling Ordination

Based on assumed affinities and ecologies of parent plants of pollen taxa recovered from the three localities and the position of taxa in ordination space, the first dimension (x-axis) appears to represent moisture levels, while the second dimension (y-axis) appears to represent disturbance (Figure 8). Using the principal of uniformitarianism, I assume that Cretaceous plants most likely had similar environmental tolerances and followed similar ecological patterns than those alive today. Therefore, members of Pteridophytina-Podocarpaceae and Pteridophytina-

Bryophyta assemblages on the left represent wet, swampy conditions. Drier, upland taxa of the

56 Bisaccate and Sciadopityspollenites-Laricoidites assemblages are to the right. Therefore, the first dimension is interpreted to represent moisture content.

Early successional Pteridophytina-Bryophyta and much of the Bisaccate assemblage are at the top of the diagram, while Sciadopityspollenites-Laricoidites and Cupressaceae-Taxaceae assemblages are towards the bottom of the diagram. This makes sense because in modern environments, Larix laricina (likely related to Laricoidites magnus) are often early to mid- successional species that colonize disturbed sites, for example, following fires (Uchytil, 1991).

Additionally, modern Cupressaceae-Taxaceae are known to form climax communities in lowland environments or on upslope environments today (Vakhrameev, 1991; Abbink et al., 2004; Dejax et al., 2007; Schrank, 2010; Galloway et al., 2013). Dicotyledonous angiosperm pollen are grouped with the Cupressaceae-Taxaceae assemblage, which is interpreted as the latest successional assemblage.

The placement of dicotyledonous angiosperm in particular was unexpected as dicotyledonous angiosperm pollen are among the highest successional taxa documented in

Figure 8. Early angiosperms are thought to have been early colonizers of disturbed riparian or other lowland environments during the Early Cretaceous, and therefore this result may reflect the small sample size, as only six dicotyledonous angiosperm pollen were recorded in this study.

In summary, the pollen and spore assemblages preserved in samples obtained from the three sections indicate that late Albian climate on Axel Heiberg and Ellesmere islands were humid and temperate. Coastal plain lowlands were likely warm-temperate and were vegetated by

57 a variety of Pteridophytina, Bryophyta, Lycopodiopsida and Cupressaceae-Taxaceae. There were likely many small lakes or swamps, areas suitable for colonization by Bryophyta and

Pteridophytina, whereas higher areas of the coastal plain, near these lakes or swamps, could support trees of the Cupressaceae-Taxaceae, Cycadopsida/Ginkgopsida and possibly early angiosperms. Highlands were likely cooler and drier and/or better drained and forests were composed of a variety of coniferous tress, including Cupressaceae-Taxaceae and Pinus/Picea parent plants, with an understory of Pteridophytina and Bryophyta. These environments may have been subjected to frequent disturbance. These plants pollen and spores likely travelled to the coast via wind-driven pollen rain and via water in rivers and streams (Faegri and Iversen,

1989; Traverse, 2007).

In general, paleoclimatic interpretations based on the Hassel Formation and flora confirms previously published high latitude late Early and early Late Cretaceous climatic inferences. In general, across this Cretaceous interval, paleoclimate reconstructions show that a humid greenhouse climate, with cool temperate conditions prevailing at higher latitudes were characteristic in the northern hemisphere (Hopkins and Balkwill, 1973; Spicer and Parrish, 1986;

Herngreen et al., 1996; Gale, 2000; Hay, 2008; Herman and Spicer, 2010; Galloway et al.,

2012). Therefore, the climate inferred from Albian to Cenomanian Christopher, Hassel and

Bastion Ridge formations supports previous interpretations of a humid, temperate environment at high latitudes in the Northern Hemisphere.

58 CONISS

Each taxa cluster identified using R-mode cluster analysis is present at all three locations,

albeit in very different relative proportions (Figures 10 to 12). This also rejects the null

hypothesis that each section contains similar palynoassemblages. I interpret hat changes in the

relative proportions of taxa clusters delineated into stratigraphically constrained zones by

CONISS may represent ecological changes associated with changing climate over time. The

relative abundance of Cupressaceae-Taxaceae pollen increases up-section at Strathcona and

Cañón fiords (zones SF-B, SF-C and CF-A), a relationship that suggests expansion of

Cupressaceae-Taxaceae communities with the addition of other conifers (Laricoidites and

Sciadopityspollenites). These other conifer types likely began to diversify in upland areas to form mixed forest communities (SF-C- and CF-B), which expanded possibly in response to fires in the region based on comparison to relative pollen abundances of these taxa in modern environments

(Ahlgren and Ahlgren, 1960; Rowe and Scotter, 1973). Cupressaceae-Taxaceae pollen increases markedly near the top of the two sections (SF-D; CF-B). This abundance is suggestive that

Cupressaceae-Taxaceae parent plants achieved maximum abundance in the lowland or upland environments in the region. The addition of pollen with affinities to the

Cycadopsida/Ginkgopsida provides further evidence that the climate was warm enough to ward off frost in the winter and that well-drained or relatively dry slope environments existed and supported these parent plants.

At Glacier Fiord, lowland environments appear to have supported a mixture of

Pteridophytina, Bryophyta and Cupressaceae-Taxaceae, with fewer inputs from upland forest taxa (e.g. bisaccate pollen) than in the other two sections. No major shifts in taxa are recorded

59 throughout the Glacier Fiord section, suggesting relative climatic stability at this locality during deposition of the Hassel Formation. Stratigraphic palynological zone GF-E is present in samples from the Bastion Ridge Formation at the Glacier Fiord section. This zone is composed of relatively higher percentages of bisaccate and Cupressaceae-Taxaceae pollen and relatively lower percentages of Pteridophytina, Bryophyta and Lycopodiopsida spores than the Hassel and

Christopher formations. Since the Bastion Ridge Formation is interpreted as a marine mudstone, this distribution pattern could be due to hydrological sorting processes acting on conifer pollen.

In particular, bisaccate-type pollen, if not pierced or broken, tends to float on water for significant distances before becoming water-logged and sinking (Holmes, 1994). Bisaccate pollen can also travel substantial distances (100s of kms) through the air prior to deposition on water or land (Holmes, 1994). Cupressaceae-Taxaceae pollen lacks the long-distance travelling abilities of bisaccate pollen due to its very different morphology (Traverse, 2007). Some question as to the preservation potential of this relatively thin-walled pollen type exists, but

Cupressaceae-Taxaceae pollen is well represented in modern marine sediments (Mudie and

McCarthy, 2006).

Thick walled spores such as those of the genera Lycopodium (Lycopodiopsida) tend to drop out of the water column near discharge sites, while lighter pollen grains can be transported further offshore (Holmes, 1994). Given that the Bastion Ridge Formation is a marine mudstone with relatively higher bisaccate and Cupressaceae-Taxaceae pollen and relatively less heavy

Pteridophytina, Bryophyta and Lycopodiopsida spores, this distribution pattern could represent an aqueous sorting effect.

60 Q-mode Cluster Analysis

Similar flora contained in samples from all three sections of Hassel Formation studied suggests that climatic and resulting ecological conditions were broadly comparable at all three locations, which are located about 400-500 km apart from each other (Figure 1). However, Q- mode cluster analysis shows that Glacier Fiord (Axel Heiberg Island) has different relative proportions of common taxa than at the Strathcona and Cañón fiords localities, Ellesmere Island

(Figure 10). This Glacier Fiord assemblage (GF assemblage) is dominated by pollen and spores attributable to Cupressaceae-Taxaceae, Pteridophytina and Bryophyta in addition to a relatively low abundance of other conifer pollen in comparison to the other two sections. The lower part of the Strathcona Fiord section also contains a distinct assemblage (SF assemblage). These lower

Strathcona Fiord samples are dominated by undifferentiated bisaccate and Pteridophytina spores, with a relatively low abundance of Cupressaceae-Taxaceae pollen. Notably, upper Strathcona

Fiord samples (greater than about 123 m) and Cañón Fiord samples (SF/CF zone) seem to show a similar palynoflora. Upper Strathcona and Cañón fiords samples contain mostly Cupressaceae-

Taxaceae, undifferentiated bisaccate pollen and spores attributable to Pteridophytina. Because samples generally plot with other samples from the same section I reject the null hypothesis that

“samples from Hassel Formation, irrespective of location contain similar palynoassemblages.”

Instead, each section contains a distinct flora.

The relatively high percentages (46%) of undifferentiated bisaccate pollen documented in lower samples of the Strathcona Fiord section indicate that the depositional environment preserved initially received substantial pollen input from an upland environment populated by a dense coniferous (Pinus/Picea?) forest. For instance, in Holocene studies of lake sediments,

61 Pinus pollen percentages of greater than 30% and greater than 5% Picea percentages are taken to represent local growth of parent plants near the site of deposition (Lisitsyna et al., 2011). Up section at Strathcona and Cañón fiords, relative abundances of undifferentiated Bisaccate pollen are lower (21%) than at lower Strathcona Fiord (46%). Cupressaceae-Taxaceae pollen are relatively higher at upper Strathcona and Cañón fiords (32% at Strathcona Fiord vs 63% at upper

Strathcona and Cañón fiords). This suggests that at upper Strathcona Fiord, the Bisaccate parent plants had dominated the lower Strathcona Fiord assemblage became reduced in number and/or that Cupressaceae-Taxaceae parent plants had expanded to become more numerous.

Paleogeographic reconstructions for the Glacier Fiord area indicate that this location was closer to the center of Sverdrup Basin, while Strathcona and Cañón fiords were nearer the

Sverdrup Basin edge (Embry, 1991). These relationships could indicate that Strathcona and

Cañón fiords localities preserve shallower water environments. Unfortunately, without the lithological logs from Strathcona and Cañón fiords I cannot observe lithological changes that may confirm these differences. Interestingly, the lower Strathcona Fiord section contains a much higher percentage of undifferentiated bisaccate pollen grains (46%). Since Strathcona Fiord is the closest of the three sections to the basin margin, proximity to hinterlands were Bisaccate pollen producing parent plants are assumed to have thrived, may explain the relative difference in this group between the two sections. However, the percentage of undifferentiated Bisaccate pollen declines (from 46% to 21%) while Cupressaceae-Taxaceae pollen increases (from 32% to

63%) higher up the Strathcona Fiord section and throughout the Cañón Fiord section. This would suggest instead that the upland forest retreated, that strata examined from the Strathcona Fiord

62 environment represent a more expansive lowland environment, or that climate moistened to

better support Cupressaceae-Taxaceae expansion in upland environments.

It is possible that hydrological sorting occurred, concentrating bisaccate pollen at lower

Strathcona Fiord or removing them at upper Strathcona Fiord. It is known that bisaccate pollen

are able to float large distances and have the potential to become over-represented in deep

marine sediments (Holmes, 1994). However, the Hassel Formation is interpreted to be a

shoreface deposit proximal to a delta or fluvial system, making this process unlikely (Balkwill

and Hopkins, 1976; Balkwill and Roy, 1977; Balkwill et al., 1982; Balkwill, 1983; Galloway et al., 2012; Pugh et al., 2014). Nonetheless, lithological information would be required to explore potential biases, and therefore these interpretations must remain speculative. Nonetheless, statistical analyses show that relative abundance data of long-ranging taxa has the potential to facilitate recognition of differences between geographically separate localities.

SUMMARY AND CONCLUSIONS

This study presents quantitative palynological data from the Hassel Formation at Glacier,

Strathcona and Cañón fiords. This also includes data from the Christopher and Bastion Ridge formations at Glacier Fiord. Multivariate statistical techniques are applied to relative abundance palynological data of long-ranging taxa to satisfy three purposes: 1. to document early angiosperm pollen in the Sverdrup Basin; 2. employ multivariate statistical analyses to delineate ecological groupings to provide insight into polar late Early and early Late Cretaceous

63 paleoclimate; 3. explore utility of quantitative palynology to better understand

palynostratigraphy.

The three purposes set out in this thesis have been satisfied: 1. Despite broadly equitable

climate and an environment thought to suit early angiosperm growing conditions, only a total of

eleven late Albian angiosperm pollen were documented from glacier, Strathcona and Cañón

fiords. This finding provides evidence to support the hypothesis of Galloway et al. (2012) that there was a delay in the migration and diversification of angiosperms from more southern localities such as Alberta into the Sverdrup Basin. This may have been the result of paleogeographic barriers such as the Arctic Ocean and the Western Interior Seaway or potentially an ecological or evolutionary effect related to early angiosperm insect-pollination strategies.

2. A combination of multidimensional scaling ordination and R-mode cluster analysis reveal that late Albian climate in the Sverdrup Basin and in particular on Axel Heiberg and

Ellesmere islands was humid and temperate. Coastal plain lowlands were likely warm temperate and were vegetated by a variety of Pteridophytina, Bryophyta and Lycopodiopsida, as well as trees with affinities to the Cupressaceae-Taxaceae. There were likely many small lakes or swamps, suitable areas for colonization by Bryophyta and Pteridophytina, while higher areas of the coastal plain near these lakes or swamps could support trees of the Cupressaceae-Taxaceae,

Cycadopsida/Ginkgopsida and possibly early angiosperms. Highlands were likely cooler and drier and/or better drained. Forests were composed of a variety of coniferous trees, including

Cupressaceae-Taxaceae and Pinus/Picea? parent plants, with an understory of Pteridophytina

64 and Bryophyta. Their pollen and spores were likely dispersed to the coast via pollen rain and in

rivers and streams.

On Ellesmere Island, at Strathcona and Cañón fiords, Cupressaceae-Taxaceae parent

plants expanded through the Albian to achieve climax communities near the top of the sections,

while disturbances such as fire may have allowed members of the Sciadopityspollenites-

Laricoidites assemblage to diversify upland forests. No stratigraphic changes were recorded from the Christopher and Hassel formations at Glacier Fiord, suggesting climatic stability at this locality while these formations were being deposited. However, the Bastion Ridge Formation contains a different palynoassemblages composed of higher percentages of bisaccate and

Cupressaceae-Taxaceae pollen and relatively lower percentages of Pteridophytina, Bryophyta and Lycopodiopsida spores than the Hassel and Christopher formations. This may be due to taphonomic biases such as an aqueous sorting effect of bisaccate pollen.

3. While climate and overall taxonomic signatures are similar between Glacier,

Strathcona and Cañón fiords, Q-mode hierarchical clustering show differences between samples from Axel Heiberg and Ellesmere islands. Glacier Fiord samples are dominated by pollen and spores attributable to Cupressaceae-Taxaceae, Pteridophytina and Bryophyta and a relatively low abundance of other conifer pollen in comparison to Strathcona and Cañón fiords. Lower

Strathcona Fiord samples are dominated by undifferentiated bisaccate and Pteridophytina spores, with a relatively low abundance of Cupressaceae-Taxaceae pollen, while upper Strathcona and

Cañón fiords samples contain mostly Cupressaceae-Taxaceae, undifferentiated bisaccate pollen and spores attributable to Pteridophytina. Lithological information from Ellesmere Island

65 sections, which is unavailable, is required to interpret differences between sites, but these differences could potentially represent climate, environment and/or preservation differences.

Nonetheless, palynological differences are documented between sites using primarily long- ranging, generic-level identifications of taxa. Therefore, I reject the null hypothesis “samples from Hassel Formation, irrespective of location contain similar palynoassemblages.”

Palynoassemblages differences between localities show that quantitative palynology show potential for refining palynostratigraphy using long-ranging, generic-level identifications.

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77 Appendix I: Glacier Fiord count data.

Sample Clusters Taxa A A A A A A A A A A A ID/Depth (meters) Cluster 1.5 4 6.5 11 17 19 25 33 36 76 79 Monocolpate undiff. 1A 0 1 0 0 0 0 0 0 0 0 0 Cicatricososporites spp. 1A 2 0 0 0 2 0 0 0 0 0 0 Deltoidospora hallii 1A 12 15 5 10 8 1 9 0 0 2 0 Murospora spp. 1A 1 2 0 0 0 0 0 0 2 0 0 Stoveresporites spp. 1A 0 1 0 0 0 0 0 0 0 0 0 Podocarpites spp. 1A 2 1 0 0 0 1 0 0 0 0 0 Cinguitriletes spp. 1B 0 0 0 0 0 0 0 0 0 0 0 Stereisporites antiquasporites 1B 16 5 10 29 15 17 17 25 19 13 9 Appendicisporites spp. 1B 0 0 0 0 0 0 0 0 0 0 0 Baculatisporites comaumensis 1B 6 4 3 3 3 2 0 1 4 0 5 Biretisporites potoniaei 1B 13 26 12 22 15 12 21 9 15 13 15 Camarozonosporites ambigens 1B 0 2 1 0 3 1 0 0 0 0 0 Cicatricosisporites spp. 1B 3 4 3 7 5 2 5 5 11 3 2 Gleicheniidites spp. 1B 9 6 10 13 9 11 11 17 18 5 17 Gleicheniidites senonicus 1B 32 35 35 19 31 38 29 27 36 19 33 Lycopodiumsporites expansus 1B 0 0 0 0 0 0 0 0 0 2 0 Vitreisporites spp. 1B 0 0 0 0 0 0 0 0 0 3 1 Tricolpate undiff. 2A 0 0 0 0 0 0 0 0 0 0 0 Cycadopites spp. 2A 3 1 1 1 3 7 10 7 5 1 10 Concavisporites juriensis 2A 0 0 0 0 0 0 0 1 0 0 0 Concavissimisporites spp. 2A 0 0 0 0 0 0 0 0 0 0 0 Gleicheniidites circinidites 2A 0 0 0 0 0 0 0 0 0 0 0 Lycopodiumsporites spp. 2A 4 1 1 0 0 0 2 1 0 0 1 Taxodiaceae hiatus 2A 120 156 170 152 155 168 149 152 156 203 182 Aequitriradites spp. 2B 0 0 0 0 0 0 0 0 0 0 0 Cingutriletes clavus 2B 31 7 9 11 5 8 8 12 11 14 13 Cicatricosisporites hallei 2B 0 0 0 0 0 0 0 0 0 0 0 Converrucosisporites spp. 2B 0 0 1 0 0 0 0 1 2 0 1 Cyathidites australis 2B 0 0 0 0 0 5 2 3 2 2 5 Cyathidites minor 2B 3 5 1 0 0 4 7 10 4 3 0 Deltoidospora spp. 2B 0 0 0 0 0 0 0 0 0 0 0 Laevigatosporites ovatus 2B 0 0 0 0 0 0 0 1 0 0 0 Osmundacidites wellmanii 2B 0 1 2 2 4 2 4 0 2 0 2 Pilosisporites spp. 2B 0 0 0 0 0 0 0 0 0 0 0 Trilobosporites marylandensis 2B 0 0 0 0 0 1 0 0 0 0 0 Undulatisporites spp. 2B 0 0 0 0 0 0 0 0 0 0 0 Verrucosisporites rotundus 2B 0 0 0 0 0 0 0 0 0 0 0 Verrucosisporites spp. 2B 1 3 0 0 0 1 0 0 0 0 1 Entylissa spp. 2B 0 0 0 0 0 0 0 0 0 0 0 Neoraistrickia spp. 2B 0 0 0 0 0 0 0 0 0 0 0 Undiff. bisaccate pollen 2B 56 51 36 39 49 40 37 30 17 21 11 Deltoidospora psilostoma 2C 2 0 1 1 2 1 0 0 1 0 1 Pilosisporites trichopapillus 2C 1 0 0 0 0 0 0 0 0 0 0 Equesetosporites type D 2C 0 0 0 0 0 0 0 0 0 0 0 Neoraistrickia truncata 2C 0 0 0 0 0 0 1 1 1 1 0 Laricoidites magnus 2C 0 0 0 0 0 0 0 0 0 0 0 Sciadopityspollenites spp. 2C 0 0 0 0 0 0 2 0 0 1 0

78 Appendix I: Glacier Fiord count data continued.

Sample Clusters Taxa A A A A A A C A A ID/Depth (meters) Cluster 99 117.5 131 143.5 286 286.2 286.5 286.6 288 Monocolpate undifferentiated 1A 0 0 0 0 0 0 0 0 0 Cicatricososporites spp. 1A 0 0 0 0 0 0 0 0 0 Deltoidospora hallii 1A 0 0 0 0 0 0 0 0 0 Murospora spp. 1A 0 0 0 0 0 0 0 0 0 Stoveresporites spp. 1A 0 0 0 0 0 0 0 0 0 Podocarpites spp. 1A 0 0 0 0 0 0 1 0 0 Cinguitriletes spp. 1B 0 0 0 0 0 0 0 0 0 Stereisporites antiquasporites 1B 10 22 20 9 15 7 21 18 7 Appendicisporites spp. 1B 1 1 0 0 0 0 1 0 0 Baculatisporites comaumensis 1B 2 3 12 4 0 0 0 0 1 Biretisporites potoniaei 1B 23 14 5 12 5 11 10 6 7 Camarozonosporites ambigens 1B 4 0 0 0 1 0 0 0 0 Cicatricosisporites spp. 1B 4 2 5 3 4 2 2 2 2 Gleicheniidites spp. 1B 22 14 18 18 6 9 8 3 14 Gleicheniidites senonicus 1B 54 42 21 61 51 41 25 31 43 Lycopodiumsporites expansus 1B 2 0 0 0 0 0 0 1 0 Vitreisporites spp. 1B 0 4 5 1 2 1 0 0 0 Tricolpate undifferentiated 2A 0 0 0 0 0 0 0 0 0 Cycadopites spp. 2A 4 7 5 5 0 0 9 7 4 Concavisporites juriensis 2A 0 0 0 0 0 2 0 0 0 Concavissimisporites spp. 2A 0 0 0 0 0 0 1 0 0 Gleicheniidites circinidites 2A 0 0 0 0 0 0 0 0 0 Lycopodiumsporites spp. 2A 0 0 0 0 0 0 0 1 0 Taxodiaceae hiatus 2A 157 160 181 115 145 156 220 182 159 Aequitriradites spp. 2B 0 0 0 0 0 0 0 0 0 Cingutriletes clavus 2B 10 25 11 7 2 3 8 5 0 Cicatricosisporites hallei 2B 0 0 0 0 0 0 0 0 0 Converrucosisporites spp. 2B 0 0 0 2 0 0 2 0 0 Cyathidites australis 2B 1 3 0 2 3 0 1 5 1 Cyathidites minor 2B 0 6 0 2 0 4 1 2 5 Deltoidospora spp. 2B 0 0 0 0 5 6 1 1 9 Laevigatosporites ovatus 2B 2 3 0 0 0 0 1 1 0 Osmundacidites wellmanii 2B 5 1 0 0 3 13 7 1 6 Pilosisporites spp. 2B 0 0 0 0 0 0 0 0 0 Trilobosporites marylandensis 2B 0 0 0 0 0 0 0 0 0 Undulatisporites spp. 2B 0 0 0 0 0 0 0 0 0 Verrucosisporites rotundus 2B 0 0 0 0 0 0 0 0 0 Verrucosisporites spp. 2B 0 0 2 4 6 4 3 1 2 Entylissa spp. 2B 0 0 0 0 0 0 0 0 0 Neoraistrickia spp. 2B 0 0 0 0 0 0 0 0 2 Undiff. bisaccate pollen 2B 28 51 38 55 41 64 25 46 64 Deltoidospora psilostoma 2C 0 0 0 0 0 0 0 0 0 Pilosisporites trichopapillus 2C 0 0 0 0 0 1 0 0 0 Equesetosporites type D 2C 0 0 0 0 0 0 1 0 0 Neoraistrickia truncata 2C 2 0 0 1 1 0 0 0 0 Laricoidites magnus 2C 0 0 0 0 0 0 2 0 0 Sciadopityspollenites spp. 2C 0 0 0 0 0 0 1 2 0

79 Appendix II: Strathcona Fiord count data. Sample Clusters Taxa C C C C C C C C C C ID/Depth (meters) Cluster 1 6 10 13 21 24 27 30 33 36 Monocolpate undifferentiated 1A 0 0 0 0 0 0 0 0 0 0 Cicatricososporites spp. 1A 0 0 0 0 0 2 0 0 0 0 Deltoidospora hallii 1A 0 0 0 0 0 0 0 0 0 0 Murospora spp. 1A 0 0 0 0 0 0 0 0 0 0 Stoveresporites spp. 1A 0 0 0 0 0 0 0 0 0 0 Podocarpites spp. 1A 0 0 0 2 0 0 0 0 0 0 Cinguitriletes spp. 1B 0 0 0 0 0 0 0 0 0 0 Stereisporites antiquasporites 1B 2 7 15 2 6 8 3 1 6 1 Appendicisporites spp. 1B 0 0 0 0 0 0 0 0 0 0 Baculatisporites comaumensis 1B 0 0 0 1 6 3 4 0 2 1 Biretisporites potoniaei 1B 3 0 5 1 6 6 6 11 11 8 Camarozonosporites ambigens 1B 0 0 0 0 0 0 0 0 0 0 Cicatricosisporites spp. 1B 2 5 0 2 0 4 0 0 2 0 Gleicheniidites spp. 1B 9 18 1 3 9 6 3 4 13 1 Gleicheniidites senonicus 1B 14 15 6 7 24 24 9 11 27 12 Lycopodiumsporites expansus 1B 0 0 0 0 0 0 0 0 0 0 Vitreisporites spp. 1B 0 0 0 0 0 0 0 0 0 0 Tricolpate undifferentiated 2A 0 0 0 0 0 0 0 0 0 0 Cycadopites spp. 2A 2 4 0 2 0 0 0 0 0 0 Concavisporites juriensis 2A 0 1 0 0 0 0 0 0 0 0 Concavissimisporites spp. 2A 0 0 0 0 0 0 0 0 0 0 Gleicheniidites circinidites 2A 0 1 0 0 0 0 0 0 0 0 Lycopodiumsporites spp. 2A 2 9 8 3 8 1 9 4 11 3 Taxodiaceae hiatus 2A 26 57 24 135 110 105 76 112 66 44 Aequitriradites spp. 2B 0 0 0 0 1 1 1 2 0 0 Cingutriletes clavus 2B 0 0 0 0 0 0 0 0 0 0 Cicatricosisporites hallei 2B 2 4 0 1 3 0 0 2 0 0 Converrucosisporites spp. 2B 0 0 4 0 0 0 0 0 0 0 Cyathidites australis 2B 2 4 0 0 0 3 2 1 5 2 Cyathidites minor 2B 8 18 0 0 7 8 2 5 9 3 Deltoidospora spp. 2B 0 0 0 0 0 3 2 0 6 4 Laevigatosporites ovatus 2B 2 3 3 0 2 1 1 2 8 2 Osmundacidites wellmanii 2B 7 16 5 6 6 1 3 4 3 2 Pilosisporites spp. 2B 0 2 0 1 4 0 2 0 0 0 Trilobosporites marylandensis 2B 0 0 0 0 0 0 0 0 0 0 Undulatisporites spp. 2B 0 4 0 0 1 0 0 0 0 0 Verrucosisporites rotundus 2B 3 3 0 0 0 0 0 0 0 0 Verrucosisporites spp. 2B 3 3 11 2 1 2 3 2 3 4 Entylissa spp. 2B 2 9 4 0 3 0 0 0 5 6 Neoraistrickia spp. 2B 0 1 0 0 0 0 0 0 0 0 Undiff. bisaccate pollen 2B 206 124 233 152 151 172 195 137 120 205 Deltoidospora psilostoma 2C 0 0 0 0 0 0 0 0 0 0 Pilosisporites trichopapillus 2C 0 0 0 0 0 0 0 0 0 0 Equesetosporites type D 2C 0 0 0 0 0 0 0 0 0 0 Neoraistrickia truncata 2C 3 0 0 0 0 0 0 0 0 0 Laricoidites magnus 2C 3 1 2 1 2 1 0 0 2 0 Sciadopityspollenites spp. 2C 3 0 1 2 8 4 3 6 4 2

80 Appendix II: Strathcona Fiord count data continued. Sample Clusters Taxa C C C C C C C C B C ID/Depth (meters) Cluster 46 52 53 57 60 65 67 70 73 76 Monocolpate undifferentiated 1A 0 0 0 0 0 0 0 0 0 0 Cicatricososporites spp. 1A 0 0 0 0 0 0 0 0 0 0 Deltoidospora hallii 1A 0 0 3 5 4 5 5 7 4 7 Murospora spp. 1A 0 0 0 0 0 0 0 0 0 0 Stoveresporites spp. 1A 0 0 0 0 0 0 0 0 0 0 Podocarpites spp. 1A 1 0 2 0 0 0 0 0 0 0 Cinguitriletes spp. 1B 0 1 0 0 0 0 0 0 0 0 Stereisporites antiquasporites 1B 0 2 0 3 2 0 0 0 1 0 Appendicisporites spp. 1B 0 0 0 0 0 0 0 0 0 0 Baculatisporites comaumensis 1B 3 4 0 0 0 0 0 0 1 0 Biretisporites potoniaei 1B 8 13 13 9 10 7 4 7 6 18 Camarozonosporites ambigens 1B 0 0 0 0 0 0 0 0 0 0 Cicatricosisporites spp. 1B 1 1 1 0 0 0 0 0 0 0 Gleicheniidites spp. 1B 9 8 11 7 5 11 10 9 13 10 Gleicheniidites senonicus 1B 13 41 38 41 38 28 34 19 16 25 Lycopodiumsporites expansus 1B 0 0 0 0 0 0 0 0 0 0 Vitreisporites spp. 1B 0 0 0 0 0 0 0 0 0 0 Tricolpate undifferentiated 2A 0 0 0 0 0 0 0 0 0 0 Cycadopites spp. 2A 0 0 0 6 8 5 2 2 1 5 Concavisporites juriensis 2A 0 0 0 0 0 0 0 0 0 0 Concavissimisporites spp. 2A 0 0 0 0 0 0 0 0 0 0 Gleicheniidites circinidites 2A 0 0 0 0 0 0 0 0 0 0 Lycopodiumsporites spp. 2A 3 4 2 2 1 2 5 2 3 2 Taxodiaceae hiatus 2A 92 61 103 70 56 86 55 106 174 151 Aequitriradites spp. 2B 1 0 0 0 0 0 0 0 0 0 Cingutriletes clavus 2B 0 2 0 0 0 0 0 0 0 0 Cicatricosisporites hallei 2B 0 0 0 0 0 0 0 0 0 0 Converrucosisporites spp. 2B 0 0 0 0 0 2 1 0 0 0 Cyathidites australis 2B 5 4 1 3 1 2 2 1 3 0 Cyathidites minor 2B 2 6 4 1 3 2 5 3 2 2 Deltoidospora spp. 2B 5 0 0 0 0 0 0 0 0 0 Laevigatosporites ovatus 2B 5 5 1 3 0 1 2 5 0 4 Osmundacidites wellmanii 2B 3 8 3 0 2 1 0 0 4 0 Pilosisporites spp. 2B 0 1 0 0 0 0 1 0 1 3 Trilobosporites marylandensis 2B 0 0 0 0 0 0 0 0 0 0 Undulatisporites spp. 2B 0 0 0 0 0 0 0 0 0 0 Verrucosisporites rotundus 2B 0 0 0 2 0 0 0 0 0 0 Verrucosisporites spp. 2B 0 4 2 0 0 0 0 0 0 0 Entylissa spp. 2B 10 1 0 4 5 5 3 3 5 0 Neoraistrickia spp. 2B 0 0 0 0 0 0 0 0 0 0 Undiff. bisaccate pollen 2B 163 136 149 151 160 152 177 150 143 108 Deltoidospora psilostoma 2C 0 0 0 0 2 5 1 0 2 1 Pilosisporites trichopapillus 2C 0 0 0 0 0 2 1 0 0 0 Equesetosporites type D 2C 0 0 0 0 0 1 0 0 0 0 Neoraistrickia truncata 2C 0 0 1 1 1 1 2 0 2 1 Laricoidites magnus 2C 0 0 0 12 6 6 2 4 2 2 Sciadopityspollenites spp. 2C 5 1 12 26 20 7 17 12 7 6

81 Appendix II: Strathcona Fiord count data continued. Sample Clusters Taxa C C C C B B B B B B ID/Depth (meters) Cluster 79 85 94 120 123 126 129 132 135 141 Monocolpate undifferentiated 1A 0 0 0 0 0 0 0 0 0 0 Cicatricososporites spp. 1A 0 0 0 0 0 0 0 0 0 1 Deltoidospora hallii 1A 10 2 4 1 0 2 1 3 4 9 Murospora spp. 1A 1 0 0 0 0 0 0 0 0 0 Stoveresporites spp. 1A 0 0 0 0 0 0 0 0 0 0 Podocarpites spp. 1A 1 0 0 0 0 0 0 0 0 0 Cinguitriletes spp. 1B 0 0 0 0 0 0 0 0 0 0 Stereisporites antiquasporites 1B 0 2 1 0 0 0 0 4 0 13 Appendicisporites spp. 1B 0 0 0 0 0 0 0 0 0 0 Baculatisporites comaumensis 1B 0 0 0 0 0 0 0 3 0 1 Biretisporites potoniaei 1B 11 10 11 8 6 9 8 4 6 8 Camarozonosporites ambigens 1B 0 0 0 0 0 0 0 0 0 0 Cicatricosisporites spp. 1B 0 0 0 0 0 0 0 0 0 1 Gleicheniidites spp. 1B 10 10 9 2 8 4 11 3 5 7 Gleicheniidites senonicus 1B 35 11 22 15 25 13 15 14 8 15 Lycopodiumsporites expansus 1B 0 0 0 0 0 0 0 0 0 0 Vitreisporites spp. 1B 0 0 0 0 0 0 0 0 0 0 Tricolpate undiffferentiated 2A 0 0 0 0 0 0 0 0 0 0 Cycadopites spp. 2A 2 16 8 9 7 3 3 16 12 14 Concavisporites juriensis 2A 0 0 0 0 0 0 0 0 0 0 Concavissimisporites spp. 2A 0 0 0 0 0 0 0 0 0 0 Gleicheniidites circinidites 2A 0 0 0 0 0 0 0 0 2 3 Lycopodiumsporites spp. 2A 3 4 0 3 3 3 1 2 1 17 Taxodiaceae hiatus 2A 124 172 175 165 200 165 194 220 145 197 Aequitriradites spp. 2B 0 0 0 0 0 0 0 0 0 0 Cingutriletes clavus 2B 0 0 0 0 0 0 0 0 0 0 Cicatricosisporites hallei 2B 0 0 0 0 1 0 0 0 0 1 Converrucosisporites spp. 2B 1 0 2 0 0 1 0 0 0 0 Cyathidites australis 2B 1 0 1 0 1 1 0 1 1 1 Cyathidites minor 2B 1 2 4 10 2 1 3 1 1 6 Deltoidospora spp. 2B 0 0 0 0 0 0 0 1 0 0 Laevigatosporites ovatus 2B 2 2 3 2 1 1 1 1 2 0 Osmundacidites wellmanii 2B 1 0 0 3 0 0 1 0 0 0 Pilosisporites spp. 2B 3 3 2 1 0 0 1 0 0 0 Trilobosporites marylandensis 2B 0 0 0 0 0 0 0 0 0 0 Undulatisporites spp. 2B 0 0 0 0 0 0 0 0 0 0 Verrucosisporites rotundus 2B 0 0 0 0 0 0 0 0 0 0 Verrucosisporites spp. 2B 1 0 0 11 3 5 3 1 0 0 Entylissa spp. 2B 1 0 5 5 2 4 0 3 2 5 Neoraistrickia spp. 2B 1 0 0 0 0 0 0 0 0 0 Undiff. bisaccate pollen 2B 137 80 95 75 115 94 60 73 128 83 Deltoidospora psilostoma 2C 1 0 1 0 0 0 0 0 0 0 Pilosisporites trichopapillus 2C 0 0 0 0 0 0 0 0 0 0 Equesetosporites type D 2C 0 0 0 0 0 0 0 0 0 0 Neoraistrickia truncata 2C 1 0 2 0 0 0 0 0 1 0 Laricoidites magnus 2C 1 0 2 0 0 0 0 1 5 2 Sciadopityspollenites spp. 2C 9 8 15 0 3 0 0 11 5 5

82 Appendix II: Strathcona Fiord count data continued. Sample Clusters Taxa B C A B ID/Depth (meters) Cluster 144 147 150 159 Monocolpate undifferentiated 1A 0 0 0 0 Cicatricososporites spp. 1A 0 0 0 0 Deltoidospora hallii 1A 3 4 6 2 Murospora spp. 1A 0 0 0 0 Stoveresporites spp. 1A 0 0 0 0 Podocarpites spp. 1A 0 0 0 0 Cinguitriletes spp. 1B 0 0 0 0 Stereisporites antiquasporites 1B 9 6 3 15 Appendicisporites spp. 1B 0 0 0 0 Baculatisporites comaumensis 1B 0 0 0 0 Biretisporites potoniaei 1B 2 10 6 6 Camarozonosporites ambigens 1B 0 0 0 0 Cicatricosisporites spp. 1B 0 2 0 1 Gleicheniidites spp. 1B 3 2 2 7 Gleicheniidites senonicus 1B 9 18 7 16 Lycopodiumsporites expansus 1B 0 0 0 0 Vitreisporites spp. 1B 0 0 0 0 Tricolpate undifferentiated 2A 0 0 0 0 Cycadopites spp. 2A 25 8 6 12 Concavisporites juriensis 2A 0 3 0 0 Concavissimisporites spp. 2A 0 3 0 0 Gleicheniidites circinidites 2A 0 1 0 1 Lycopodiumsporites spp. 2A 5 13 1 11 Taxodiaceae hiatus 2A 180 265 235 200 Aequitriradites spp. 2B 0 0 0 0 Cingutriletes clavus 2B 0 0 0 0 Cicatricosisporites hallei 2B 1 0 2 1 Converrucosisporites spp. 2B 0 0 0 0 Cyathidites australis 2B 2 2 1 0 Cyathidites minor 2B 2 4 2 1 Deltoidospora spp. 2B 0 0 0 0 Laevigatosporites ovatus 2B 0 0 0 0 Osmundacidites wellmanii 2B 0 0 0 0 Pilosisporites spp. 2B 0 0 0 0 Trilobosporites marylandensis 2B 0 0 1 0 Undulatisporites spp. 2B 0 0 0 0 Verrucosisporites rotundus 2B 0 0 0 0 Verrucosisporites spp. 2B 0 0 0 1 Entylissa spp. 2B 2 2 0 0 Neoraistrickia spp. 2B 0 0 1 0 Undiff. bisaccate pollen 2B 60 76 87 69 Deltoidospora psilostoma 2C 0 0 1 0 Pilosisporites trichopapillus 2C 0 0 0 0 Equesetosporites type D 2C 0 0 0 0 Neoraistrickia truncata 2C 0 0 0 0 Laricoidites magnus 2C 1 0 0 0 Sciadopityspollenites spp. 2C 2 6 1 4

83 Appendix III: Cañón Fiord count data. Sample Clusters Taxa A B B B B B B B B B ID/Depth (meters) Clusters 4 9 11 15 20 35 38 41 44 50 Monocolpate undifferentiated 1A 0 0 0 0 0 0 0 0 0 0 Cicatricososporites spp. 1A 0 1 0 0 1 0 0 0 0 0 Deltoidospora hallii 1A 4 1 0 4 3 0 2 0 2 0 Murospora spp. 1A 0 0 0 0 0 0 0 0 0 0 Stoveresporites spp. 1A 0 0 0 0 0 0 0 0 0 0 Podocarpites spp. 1A 0 0 0 0 0 0 0 0 0 0 Cinguitriletes spp. 1B 0 0 0 0 0 0 0 0 0 0 Stereisporites antiquasporites 1B 118 33 10 28 15 4 1 3 13 6 Appendicisporites spp. 1B 0 0 0 0 0 0 0 0 0 0 Baculatisporites comaumensis 1B 2 0 0 0 0 0 0 0 0 0 Biretisporites potoniaei 1B 10 9 3 0 0 0 1 7 4 1 Camarozonosporites ambigens 1B 0 0 0 0 0 0 0 0 0 0 Cicatricosisporites spp. 1B 0 0 1 0 0 0 0 1 0 1 Gleicheniidites spp. 1B 30 9 12 4 2 5 4 0 3 1 Gleicheniidites senonicus 1B 47 32 23 12 8 9 8 13 11 0 Lycopodiumsporites expansus 1B 0 0 0 0 0 0 0 0 0 0 Vitreisporites spp. 1B 0 0 0 0 0 0 0 0 0 0 Tricolpate undifferentiated 2A 0 0 0 1 2 2 0 1 0 0 Cycadopites spp. 2A 4 2 1 2 1 0 4 1 0 0 Concavisporites juriensis 2A 0 0 0 0 0 0 0 0 0 0 Concavissimisporites spp. 2A 0 0 0 0 0 0 0 0 0 0 Gleicheniidites circinidites 2A 0 0 0 0 0 0 0 0 0 0 Lycopodiumsporites spp. 2A 2 1 2 4 23 1 3 1 0 0 Taxodiaceae hiatus 2A 92 175 230 208 198 242 241 230 235 265 Aequitriradites spp. 2B 0 0 0 0 0 0 0 0 0 0 Cingutriletes clavus 2B 0 0 0 0 0 0 0 0 0 0 Cicatricosisporites hallei 2B 0 0 1 1 0 0 0 0 0 2 Converrucosisporites spp. 2B 0 1 0 0 0 0 0 1 0 0 Cyathidites australis 2B 0 0 3 0 0 1 0 0 0 0 Cyathidites minor 2B 1 1 0 2 2 1 0 1 0 0 Deltoidospora spp. 2B 0 0 0 0 0 0 0 0 0 0 Laevigatosporites ovatus 2B 0 0 1 1 0 0 0 1 1 0 Osmundacidites wellmanii 2B 0 0 2 0 0 0 0 1 0 1 Pilosisporites spp. 2B 0 0 0 0 0 0 0 0 0 0 Trilobosporites marylandensis 2B 0 0 0 0 0 0 0 0 0 0 Undulatisporites spp. 2B 0 0 0 0 0 0 0 0 0 0 Verrucosisporites rotundus 2B 0 0 0 0 0 0 0 0 0 0 Verrucosisporites spp. 2B 0 1 0 1 0 0 0 0 0 1 Entylissa spp. 2B 0 0 0 0 0 0 0 0 0 0 Neoraistrickia spp. 2B 0 0 0 0 0 0 0 0 0 0 Undiff. bisaccate pollen 2B 12 30 72 30 51 38 32 53 35 20 Deltoidospora psilostoma 2C 0 0 1 0 0 0 0 0 0 0 Pilosisporites trichopapillus 2C 0 0 0 0 0 0 0 1 0 0 Equesetosporites type D 2C 0 0 0 0 0 0 0 0 0 0 Neoraistrickia truncata 2C 0 0 1 0 0 0 0 0 0 1 Laricoidites magnus 2C 0 0 0 0 0 0 3 0 0 1 Sciadopityspollenites spp. 2C 0 0 1 1 0 0 1 0 1 0

84