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Accepted Manuscript
Proceedings of the Yorkshire Geological
Society
Cretaceous Oceanic Anoxic Event 2 in eastern England:
further palynological and geochemical data from Melton Ross
Paul Dodsworth, James S. Eldrett & Malcolm B. Hart
DOI: https://doi.org/10.1144/pygs2019-017
Received 30 October 2019
Revised 20 May 2020
Accepted 20 May 2020
© 2020 The Author(s). This is an Open Access article distributed under the terms of the Creative
Commons Attribution 4.0 License (http://creativecommons.org/licenses/by/4.0/). Published by The Geological Society of London for the Yorkshire Geological Society. Publishing disclaimer: www.geolsoc.org.uk/pub_ethics
Supplementary material at https://doi.org/10.6084/m9.figshare.c.4987205
When citing this article please include the DOI provided above.
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Cretaceous Oceanic Anoxic Event 2 in eastern England: further palynological and geochemical
data from Melton Ross
Paul Dodsworth1, James S. Eldrett2 & Malcolm B. Hart3
1StrataSolve Ltd, 15 Francis Road, Stockton Heath, Warrington WA4 6EB, U.K.
2Shell International Exploration & Production B.V., Lange Kleiweg 40, 2288 GK Rijswijk,
Netherlands.
3School of Geography, Earth & Environmental Sciences, Plymouth University, Drake Circus,
Plymouth PL4 8AA, U.K.
Correspondence: Paul Dodsworth ([email protected])
Abstract. The lowermost 1.45 m of the Welton Chalk Formation, including the regional sedimentary
record of Oceanic Anoxic Event 2 (OAE-2), has been sampled at Melton Ross Quarry in eastern
England, U.K. The section is investigated for organic geochemistry and stable isotopes for the first
time, while a detailed palynological study follows previously published preliminary results. It
comprises a condensed interval that spans the Cenomanian–Turonian Stage boundary. A locally
preserved, lower ‗anomalous‘ succession (Beds I–VII) and a ‗Central Limestone‘ (Bed A) are shown
to correlate respectively with the pre-Plenus sequence and Plenus Bed at Misburg and Wunstorf in the
Lower Saxony Basin (LSB), NW Germany. They are overlain by a succession of variegated marls
(Bed B to Bed H), including the Black Band (Beds C–E), that can be correlated across eastern
England. Based on a carbon isotope (δ13C) profile and dinoflagellate cyst and acritarch bio-event
correlation, Beds B–H appear to be a highly attenuated post-Plenus equivalent of the LSB succession,
including part of the ‗Fish Shale‘. The δ13C profile shows possible ‗precursor‘ / ‗build-up‘ events in
the lower succession at Melton Ross, with the main OAE-2 δ13C excursion occurring in the Central
Limestone and overlying Beds B–H. The darker coloured marls from the Black Band and Bed G
contain 1.43–3.47% total organic carbon (TOC), hydrogen index values of 78–203 mg HC/g TOC and
oxygen index values of 15–26 mg CO2/g TOC, indicating type III and type II–III organic matter, of
mixed terrigenous and marine algal sources. The corresponding palynological assemblages are
dominated by marine dinoflagellate cysts, comprising mainly gonyaulacoid taxa, with subordinate
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terrigenous miospores, mainly gymnosperm bisaccate pollen, consistent with a distal marine setting.
The interbedded lighter-coloured marls contain less than 0.4% TOC and lower proportions of
miospores and peridinioid dinoflagellate cysts compared with the darker layers. This is suggestive of
moderately raised levels of productivity during deposition of the darker layers, possibly related to
greater nutrient availability from land-derived sources. The occurrence of the peridinioid taxa
Eurydinium saxoniense and Bosedinia spp., together with higher proportions of prasinophyte
phycomata in the darker layers, may also point to stimulation of organic-walled phytoplankton
productivity by reduced nitrogen chemo-species encroaching the photic zone, possibly by expansion
of an oxygen-minimum zone. Exceptionally high concentrations of palynomorphs (in the tens of
thousands to lower hundreds of thousands per gramme range) in the darker layers at Melton Ross and
eight other eastern England localities is consistent with increased quality of sea floor preservation in a
low oxygen environment, coupled with a high degree of stratigraphic condensation. Two new
dinoflagellate cyst species are described from Melton Ross, Canninginopsis? lindseyensis sp. nov. and
Trithyrodinium maculatum sp. nov., along with two taxa described in open nomenclature.
Supplementary material: One pdf file, with detailed sample positions and descriptions, tables of
supporting information (also available in Excel format), quarry photographs and a palynological
distribution chart, is available at
The Late Cretaceous Epoch was characterized by sustained warm climate, resulting in high eustatic
sea levels (Miller et al. 2005; Haq & Huber 2016; Hay et al. 2018). Numerous epicontinental seaways
became established, submerging large areas of Western Europe (e.g. Gale 1995) and the Western
Interior of North America (Kauffman & Caldwell 1993). Major global perturbations in the carbon
cycle occurred, termed Oceanic Anoxic Events, the most prominent spanning the Cenomanian–
Turonian Stage boundary (CTB, 93.9 Ma; International Commission on Stratigraphy 2019), named
OAE-2 (Schlanger & Jenkins 1976; Schlanger et al. 1987) and lasting for up to ca. 900 kyr (Eldrett et
al. 2015a; Gangl et al. 2019). This interval was marked by a globally recognized positive carbon
isotope (δ13C) excursion, reflecting the widespread sequestration of δ12C-enriched organic matter in
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marine sediments under global anoxic conditions (Jenkyns 2010 and references therein). However, the
deposition of dark-coloured, organic-rich, fine-grained sediments (‗black shales‘) varied both
temporally and spatially, being modulated and ultimately dependent on local and regional processes
(basin restriction, water stratification, bottom-currents, sediment input) in addition to global
phenomena (Large Igneous Province activity, sea level change, orbital forcing; e.g. Ernst & Youbi
2017; Clarkson et al. 2018; Minisini et al. 2018). A global mass extinction / turnover bio-event
occurred around the CTB and is probably associated with OAE-2 (Raup & Sepkoski 1982, 1984;
Kauffman 1984; Milne et al. 1985; Hart 2005, 2019).
In the North Sea Basin and adjacent outcrop areas in NW Germany and eastern England, deposition of
dark coloured mudstones interbedded with light coloured mudstones and limestones commenced
during the Late Cenomanian, and continued into the Early Turonian in some areas, including
Wunstorf and Misburg in the Lower Saxony Basin, NW Germany (Fig. 1). At these locations, the
CTB succession is 12–30 m thick (Ernst et al. 1983; Hilbrecht 1986). The dark mudstone layers form
a distinctive interval in predominantly whitish limestone chalk successions. In eastern England (Fig.
2), a highly condensed (relative to NW Germany) ‗Black Band‘ is developed in Yorkshire (0.3–0.7 m
thick at Flixton, East Knapton, Bishop Wilton and Market Weighton; Jeans et al. 1991; Dodsworth
1996) and northern Lincolnshire (0.1–0.2 m thick at South Ferriby, Elsham, Melton Ross, Bigby and
Caistor). Jenkyns (1985) envisaged a pelagic shelf depositional setting for the Black Band and
adjacent strata in a relatively shallow (several hundred metres) epicontinental sea. The Black Band
wedges out to the south of Louth (Hart et al. 1991). It appears to represent the ‗feather-edge‘ of OAE-
2, which dies out when traced toward a palaeo-high that appears to have been located in the region of
the Wash (Hart et al. 1991). Schlanger et al. (1987) suggested that it may have been deposited near
the upper limit of an oxygen-minimum zone that lapped onto the continental shelf.
The Cenomanian and Turonian stages probably record the highest sea levels within the Cretaceous
Period, with maximum eustatic sea levels being reached during the Early Turonian (Haq et al. 1988;
Sahagian et al. 1996; Miller et al. 2003, 2005). They were probably the time of minimal continental
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relief during the Phanerozoic, and the time of minimal detrital sediment delivery to the ocean (Hay et
al. 2018). The CTB interval was also a time of extreme warmth with mid-latitude sea-surface
temperature possibly exceeding 35°C (Huber et al. 2002; Voigt et al. 2004; Forster et al. 2007;
Robinson et al. 2019). This warmth was likely associated with a more stratified water column,
resulting in poor atmosphere–ocean gas exchange, oxygen-deficient photic zone waters, including
euxinic conditions in some regions, and enhanced organic matter preservation at the sediment–water
interface (Sinninghe Damsté & Köster 1998; Monteiro et al. 2012). Rhythmic bedding in CTB
successions has been linked to orbital-forcing (e.g. Arthur et al. 1986; Eldrett et al. 2015b; Boulila et
al. 2020), which in some regions, including NW Germany, may have caused an intensified
hydrological cycle during warmer/wetter periods (van Helmond et al. 2015; Charbonnier et al. 2018;
Gharaie & Kalanat 2018).
The early stages of OAE-2 were characterised by a relatively extended interval (ca. 150–200 kyr;
Clarkson et al. 2018) of ca. 3–5°C cooling of sea surface temperatures in the proto-Atlantic and the
European shelf (Forster et al. 2007; Pearce et al. 2009; Sinninghe Damsté et al. 2010; Jarvis et al.
2011; van Helmond et al. 2014, 2015). Boreal Realm fauna, including the belemnite Praeactinocamax
plenus (Blainville), migrated southwards across Europe (Jefferies 1962, 1963; Gale & Christensen
1996; Marcinowski et al. 1996; Košták et al. 2004). The interval was termed the ‗Plenus Cold Event‘
(PCE) by Gale & Christensen (1996) and probably marks a shift towards improved oxygenation of
bottom waters during OAE-2 (Forster et al. 2007; Eldrett et al. 2014, 2017; van Helmond et al. 2014),
possibly on a global scale (Clarkson et al. 2018; O‘Connor et al. 2020), and relatively drier climates
(Heimhofer et al. 2018; Gharaie & Kalanat 2018). Across regions of Europe that record organic-rich
sedimentation during OAE-2, including NW Germany and SE France (e.g. Pont d'Issole, Fig. 1),
‗black shale‘ deposition temporarily gave way in some areas to more oxygenated sediments during the
PCE, and a resumption of deposition of relatively thick biogenic pelagic limestones (Wiese et al.
2009; Jarvis et al. 2011; Grosheny et al. 2017; Jenkyns et al. 2017). In NW Germany, a conspicuous
tripartite limestone bed, called the Plenus Bed or Plenus Bank because it yields the eponymous
belemnite, is developed at this level (Ernst et al. 1984; Hilbrecht & Dahmer 1994), separating
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underlying pre-Plenus and overlying post-Plenus successions of dark coloured mudstones interbedded
with light coloured mudstones and limestones.
Wood & Mortimore (1995) and Wood et al. (1997) reported the CTB succession from Melton Ross
Quarry in Lincolnshire (Fig. 2). Temporary deep excavations in the late 1990s (Fig. 3) exposed a
section hitherto unreported in eastern England, presumed eroded by unconformity elsewhere, that
appears to correlate with the lowermost 2–3 m of the NW Germany CTB succession, including the
pre-Plenus interval and the Plenus Bed. These deposits are overlain by the regionally correlative Bed
B, Black Band and Beds F–H (Fig. 4).
In a recent review of the Black Band, Hart (2019) discussed advances in foraminiferal knowledge a
quarter of a century after previous publications on their distribution across the CTB in eastern
England (Hart & Bigg 1981; Hart et al. 1993). The present paper follows up on this work, reviewing
advances in palaeoenvironmental knowledge and biostratigraphic dating using organic-walled
phytoplankton since this was last discussed for eastern England in the publications of Dodsworth
(1996, 2000) and Wood et al. (1997). Previous analysis of palynological recovery from the Black
Band at South Ferriby and Flixton (Hart et al. 1993; Dodsworth 1996) has revealed exceptionally high
concentrations of dinoflagellate cysts, in the thousands to lower hundreds of thousands per gramme
range. Hart & Koutsoukos (2015) recommended further investigation of whether the abundance of
dinoflagellate cysts is a function of their increased productivity under eutrophic conditions, increased
quality of sea floor preservation in a low oxygen environment, or a normal level of organic
productivity accentuated by a loss of biogenic carbonate sediment (including planktonic foraminifera
and calcareous nannofossils) during the CTB mass extinction (Jarvis et al. 1988a; Lamolda et al.
1994; Paul & Mitchell 1994; Hart 1996). This paper provides the first detailed palynological data
from Melton Ross and attempts to discriminate the relative contribution of these factors. A taxonomic
review of new and problematic dinoflagellate cysts is included. Other marine palynomorphs are
documented, including prasinophyte phycomata and acanthomorph acritarchs, along with land-
derived (terrigenous) pollen and spores. For regional comparison, summary palynological data are
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published for eight correlative eastern England localities. From south to north, these are Caistor,
Bigby, South Ferriby, Market Weighton, Bishop Wilton, East Knapton, Flixton and Speeton (Fig. 2).
Total organic carbon, Rock-Eval pyrolysis and stable isotope data are presented for the first time from
Melton Ross. The integrated bio- and chemostratigraphy of the section is assessed in an inter-regional
context.
In this paper, chronostratigraphic substages and their corresponding ages are treated as formal units,
using initial upper case letters, consistent with the usual practice, although not all have been ratified.
The base of the Cenomanian Stage and Lower Cenomanian Substage have a ratified Global boundary
Stratotype Section and Point (GSSP) at Mont Risou in southern France (Kennedy et al. 2004). The
base of the Turonian Stage and Lower Turonian Substage have a ratified GSSP near Pueblo, Colorado
(Fig. 1; Kennedy et al. 2005). The base of a Middle Cenomanian Substage has a proposed GSSP in
Sussex, southern England (Tröger et al. 1996), and that of a Middle Turonian Substage is proposed
near Pueblo (Bengtson et al. 1996; Kennedy et al. 2000; Dodsworth & Eldrett 2019). Tröger et al.
(1996) recommended southern France as a suitable region for an Upper Cenomanian Substage GSSP,
though they did not propose a site. Replacement of Acanthoceras ammonites by the genus
Calycoceras (Hancock 1991), which can also be correlated using carbon isotopes (Kennedy & Gale
2006), provides a possible datum for the base of an Upper Cenomanian Substage.
Lithostratigraphy
The Late Cretaceous Epoch in England is represented by the bio-micritic, white limestones of the
Chalk Group (Fig. 2), which Wood & Smith (1978) grouped into three major faunal and depositional
provinces: a Northern Province (investigated here) which links eastern England to the north of the
Wash with contemporaneous North Sea and NW German successions; a Southern Province linking
southern England and northern France (Anglo-Paris Basin); and a Transitional Province in the
Chilterns and East Anglia. In eastern England, chalk bedding is developed on a decimetre scale with
more clay-rich marls forming thin (<5 cm) inter-beds (e.g. Hancock 1976; Jeans 1980). From Louth
northward in Lincolnshire and Yorkshire, the lowermost ca. 0.5 to 1.5 m of the Welton Chalk
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Formation (Wood & Smith 1978) is characterised by an atypically thick succession of variegated
marls, including the organic-rich Black Band (Fig. 4).
Hopson (2005) gave a comprehensive review of the historical naming of the lowermost part of the
Welton Chalk Formation, along with evidence for its stratigraphic position, based on
chemostratigraphy (stable carbon isotopes: Schlanger et al. 1987; Hart et al. 1991; Gale et al. 1993;
Wood & Mortimore 1995; and subsequently, Clarkson et al. 2018) and macrofossils (e.g. Jefferies
1963; Whitham 1991; Gaunt et al. 1992; Wood et al. 1997). Hopson (2005) proposed using the term
Plenus Marl Member for marl and limestone beds between the base of the Welton Chalk Formation
and the base of the Black Band, on the grounds of their probable correlation with the Plenus Marls of
the Anglo-Paris Basin. He also proposed using the term Black Band Member for marls and limestones
between the base of the Black Band and the base of the Buckton Member (Mitchell 2000), on the
grounds of these deposits probably being stratigraphically higher than the Plenus Marls, equivalent to
part of the Melbourn Rock Member / Ballard Cliff Member in southern England. However,
subsequent workers have retained one unit, the Flixton Member of Jeans (1980), for both intervals
(Hart 2019; Mitchell 2019). In Table 1, the subdivision of the Flixton Member into two sub-members
(Brett et al. 2018) is suggested, based on the definitions of Hopson (2005).
In the Anglo-Paris Basin, Jefferies (1963) labelled the eight correlative beds of a ‗standard‘ Plenus
Marls succession as Beds 1–8 in ascending order. Any strata that could not be correlated with these
beds were labelled locally with lower case Roman numerals. Jefferies (1963) could not trace Beds 1–8
farther north than Marham in Norfolk (Fig. 2), though subsequent work (Voigt et al. 2006) tentatively
identified the uppermost units (Bed 7 and Bed 8) at Heacham and Barret Ringstead, near Hunstanton.
Jefferies (1963) investigated one locality from Lincolnshire (South Ferriby) and another from
Yorkshire (Speeton), with the Black Band assigned to ‗bed i‘ at Speeton and ‗bed iii‘ at South Ferriby.
Hart et al. (1991, 1993) applied Jefferies‘ (1963) informal bed names at South Ferriby. Subsequently,
individual beds within the Flixton Member have been assigned different names by different authors
during research published in the 1990s (Table 1). Most of these schemes use ascending numbers
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comparable to the Anglo-Paris Basin succession whilst not intending to imply lithostratigraphic
correlation with southern England beds of the same number. To avoid confusion, Dodsworth (1996)
alternatively proposed using letters rather than numbers for the eastern England succession, and
erected a regional lithostratigraphic scheme of comparable resolution to previous schemes.
In hydrocarbon wells from the central North Sea Basin, e.g. well 47/10-1 (Fig. 2), the distinctive
black, grey and green lithologies described from the CTB interval (Rhys 1974; Burnhill & Ramsay
1981) are comparable to those from the lowermost part of the Welton Chalk Formation of onshore
eastern England, and contain comparable foraminiferal assemblages (Burnhill & Ramsay 1981;
Crittenden et al. 1991) and palynological assemblages (Marshall & Batten 1988; Dodsworth 1996).
The term ‗Plenus Marl Formation‘ was initially applied to the corresponding high gamma log-
response unit in U.K. sector North Sea wells by Deegan & Scull (1977), but this has since been
replaced by ‗Black Band Bed‘ (Johnson & Lott 1993; Surlyk et al. 2003; van der Molen & Wong
2007) to reflect lithologies and stratigraphy different from that of the Plenus Marls in the Anglo-Paris
Basin. Farther north, in the Norwegian sector, e.g. well 35/6-2 S (Fig. 1), the term Blodøks Formation
(of Isaksen & Tonstad 1989) is applied to the correlative interval developed within the siliciclastic
Shetland Group. The Blodøks Formation is usually a few metres thick and rarely exceeds 20 m
(Gradstein & Waters 2016).
In this paper, the lithostratigraphy of Dodsworth (1996) is retained (Fig. 4). Higher resolution
subdivision around the Central Limestone (of Wood et al. 1997; Bed A), as described by Wood et al.
(1997) and Hildreth (1999) at Melton Ross Site 2 and some other northern Lincolnshire quarries,
including Bigby, are treated here as subdivisions of Beds A and B (Table 1). In Yorkshire, Beds A
and B may pinch out at some localities, e.g. Bishop Wilton and East Knapton (Dodsworth 1996), and
Speeton (Jefferies 1963; Mitchell et al. 1996; Mitchell 2000), with the Black Band at the base of the
Welton Chalk Formation or a short distance above it, i.e. above an attenuated, 2–12 cm thick Bed B.
The local Beds I–VII of Wood et al. (1997) are applied here to the lower succession at Melton Ross.
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Previous research
Wood & Mortimore (1995) made an initial description of a relatively expanded stratigraphic
succession at Melton Ross Quarry (Site 1; Fig. 3), which exposed a section of marls, hitherto
unreported in eastern England, above the base of the Welton Chalk Formation. The marls are
presumed to have been eroded by unconformity elsewhere. Wiese et al. (2009) suggested this lower
‗anomalous‘ section may have been fortuitously preserved in a down-faulted block, though there is no
regional geophysical evidence of such a structure.
Wood et al. (1997) described three further Melton Ross excavations made during 1995–1996 (sites 2–
4; Fig. 3), which contain a thicker lower succession than Site 1, and categorised the localised deposits
into seven units, Beds I–VII. They expressed uncertainty about how Beds III–VI correlate with Site 1.
At the time of writing (2019), sites 1–4 are buried approximately 10 m below the restored surface of
the quarry.
In addition, Wood et al. (1997) collected their own set of 27 samples from Site 2 (labelled CJW-1 to -
26) and presented Rare Earth Element data, analysis of clay mineralogy and preliminary palynological
comments from those samples. They reported shale-normalised Rare Earth Element profiles to be sub-
horizontal with no Europium anomaly, suggesting to them a detrital rather than volcanogenic
provenance for the CTB succession (Wood et al. 1997, fig. 5). They further reported the clay
mineralogy to comprise a mixture of illite, kaolinite and smectite at the base of the succession, but
becoming predominantly smectite upwards with the loss of kaolinite in the regionally correlative
upper succession (Wood et al. 1997, fig. 4). They interpreted this change in mineralogy as probably
suggesting either deepening or increasing distance from shorelines through the CTB interval (Wood et
al. 1997, p. 339).
Wood et al. (1997, p. 342, fig. 3) recovered Praeactinocamax plenus from Site 1 at Melton Ross, in
silty lithologies immediately above the Central Limestone (Fig. 4). They considered the latter to
correlate with the Plenus Bed in NW Germany. Wood & Mortimore (1995) and Wood et al. (1997)
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did not publish logs of the chalk section above the variegated marls (Bed B to Bed H) at Melton Ross,
i.e. from the upper part of the Flixton Member. This section is logged in the present study (Fig. 4) and
reveals a thickness of chalk (0.65 m) between the top of the variegated marls and ‗Adrian‘s pair of
marls‘ (of Mortimore 2014), immediately below the base of the Buckton Member, comparable to that
at nearby South Ferriby and Caistor (Hart et al. 1991, fig. 2; Mortimore 2014, fig. 4.23a; Hart 2019,
fig. 4).
Previous palynological work
There have been several previous palynological investigations of the lowermost part of the Welton
Chalk Formation. R.J. Davey, in Hart & Bigg (1981), reported dinoflagellate cysts to be the dominant
palynomorphs at Elsham (Fig. 2), with subordinate bisaccate pollen and very rare spores. Hart & Bigg
(1981) suggested that marine algae were probably the main source of the abundant organic matter in
the Black Band. They also noted that adjacent marls and chalks at Elsham were palynologically
barren, probably a function of unfavourable lithologies for the preservation of organic-walled
microfossils. A species list and distribution chart were not provided. Marshall (1983, p.172–174) and
Marshall & Batten (1988) undertook detailed analysis of six South Ferriby samples. Cyclonephelium
compactum–membraniphorum was reported to be the dominant dinoflagellate cyst in the Black Band,
with sparse recovery of palynomorphs from marls immediately below and above (Beds B and F), and
palynologically barren samples from adjacent chalks. Microplankton compose 76% of an assemblage
from the lower part of the Black Band (Bed C) and 89% from the upper part (Bed E). The
Cenomanian marker taxon Litosphaeridium siphoniphorum was not recorded in any of the samples.
Duane (1992, p. 277–292; in Hart et al. 1993) provided palynological data for four productive
samples from the east wall at South Ferriby quarry (Beds C to G); adjacent chalks and marls from ca.
75 cm above and 65 cm below were palynologically barren. Cyclonephelium compactum–
membraniphorum, along with Eurydinium saxoniense, are dominant in the dinoflagellate cyst
assemblages of the productive samples, while terrigenous bisaccate pollen and spores compose < 5%
of the total palynological assemblages. Exceptionally high concentrations of dinoflagellate cysts
(thousands per gramme), relative to Middle Cenomanian chalks and marls of southern England
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(hundreds per gramme; Paul et al. 1994), were reported. Duane (1992; in Hart et al. 1993) did not
record Litosphaeridium siphoniphorum in any of the South Ferriby samples.
Dodsworth (1996) published palynological data from three samples of the Black Band from the south
wall at South Ferriby quarry (recovery from adjacent marl samples was sparse), six samples from the
Black Band at Flixton (splits of the same samples analysed for geochemistry by Jeans et al. 1991),
two samples from Bed G at Market Weighton, and commented on the palynology of the Black Band
at Caistor, Bigby, Market Weighton, Bishop Wilton, East Knapton and Speeton. As with previous
investigations, Litosphaeridium siphoniphorum was not recorded in any of the samples. However,
taxa whose last occurrences Marshall & Batten (1988) calibrated approximately to an influx of the
latest Cenomanian zonal ammonite Neocardioceras juddii Barrois & Guerne in NW Germany
(Hilbrecht 1986), namely Adnatosphaeridium tutulosum (in Bed C at Flixton) and Carpodinium
obliquicostatum (in Bed C at South Ferriby, and Beds C and E at Flixton), were recorded and
illustrated. Batten, in Wood et al. (1997), studied two samples from the Black Band at Melton Ross
(one each from Bed C and Bed E) and confirmed an absence of L. siphoniphorum. However, analysis
of five samples from local Beds II–VI at Site 2 revealed the first records of this taxon in the Northern
Province, where it is common in occurrence. Dodsworth (2000, fig. 12), provided preliminary marker
taxa distribution data from the samples documented more fully here (Fig. 4), corroborating Batten‘s
record of common L. siphoniphorum in local Beds I–?III at Melton Ross (Site 1), and indicating the
presence of A. tutulosum and C. obliquicostatum higher in the succession (Site 4), from Beds C–F and
Bed C, respectively.
Material and methods
Twenty-seven channelled (composite) rock samples (MR97 series) were collected from the top of the
Ferriby Chalk Formation and lowermost 1.45 m of the Welton Chalk Formation in Melton Ross
Quarry by one of us (PD) on 24th February 1997 and are documented in the present paper. Eleven of
our samples (MR97-23 to -14) were collected from Site 1 of Wood & Mortimore (1995). Sites 2 and 3
and the lower part of Site 4 of Wood et al. (1997) were submerged below the quarry‘s water table in
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February 1997 and were unavailable for sampling. The upper part of Site 4 was accessible and 16 of
our samples (MR97-13 to -1) were collected from the relatively freshly-exposed, regionally
correlative beds there. The thickness of channelled samples varies from 3 mm to 230 mm, depending
on the thickness of rock layers (Fig. 4; see the Supplementary Appendix for details of the samples).
The samples were prepared and analysed for organic geochemistry, palynology, and carbonate stable
isotopes. In the following text, the sample series prefix is not written out in full for each sample; thus,
sample MR97-1 is shortened to sample -1.
Organic geochemistry
All samples were analysed for Total Organic Carbon (TOC). Samples with > 0.25% TOC (-1 to -11.5,
and -22) were also analysed for Rock-Eval pyrolysis. Analyses were undertaken in the laboratory of
Applied Petroleum Technology (APT) AS, Oslo, Norway. All procedures followed Weiss et al.
(2000). For TOC, a Leco SC-632 instrument was used. Diluted hydrochloric acid (HCl) was added to
the crushed rock samples to remove carbonate. The samples were then introduced into the Leco
combustion oven, and the amount of carbon in the sample was measured as carbon dioxide (CO2) by
an IR-detector. For Rock-Eval pyrolysis, a HAWK instrument was used. Jet-Rock 1 was run as every
tenth sample and checked against the acceptable range given in Weiss et al. (2000). The temperature
programme was a five minutes purge before pyrolysis: 300°C (three minutes) plus 25°C per minute
until 650°C was reached.
Palynological processing
Five grammes (dark coloured lithologies) or ten grammes (light coloured lithologies) of crushed,
dried material from samples -1 to -22 were dissolved in hydrochloric acid (35% HCl) and
hydrofluoric acid (40% HF) in order to remove carbonate and silicate minerals, respectively.
Preparations were sieved with a 10 μm mesh. Kerogen slides were prepared at this stage.
Palynomorphs (illustrated in Figs. 5–7), brown and black wood fragments (vitrinite and inertinite
phytoclasts) and clumps of granular amorphous organic matter (AOM) tend to occur in comparable
proportions in the > 10 μm kerogen from the < 1% TOC samples at Melton Ross (Table 2; Fig. 8). To
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improve remaining residues for the counting of palynomorphs, many of these preparations were
cleaned by treatment with a ‗nitric wash‘, i.e. one minute of oxidation with nitric acid (70% HNO3),
or a rinse with potassium hydroxide solution (2% KOH). In the > 10 μm kerogen fraction from the >
1% TOC samples, dark coloured, clumped AOM is dominant (Table 2; Fig. 8). To liberate
palynomorphs from the AOM, extended oxidation was given with: 1, nitric acid (18–24 hours),
followed by one to two minutes in an ultrasonic bath with a 2% KOH solution that was supersaturated
with potassium permanganate (KMnO4; samples -8, -8.5, -10, -11, -11.5); 2, Schulze‘s solution (nitric
acid supersaturated with potassium chlorate, KClO3), followed by one subsequent rinse with a 2%
KOH solution (sample -2). The former technique was found to give better results than the latter in
terms of palynomorph preservation and in not selectively destroying gonyaulacoid dinoflagellate cysts
(cf. Dodsworth 1995; 2004a, fig. 4; Dodsworth & Eldrett 2019). All oxidised preparations were
stained with Safranin O solution (red stain). Full laboratory processing records are available in
Supplementary Table A.
Where palynological recovery permitted, approximately equal portions of quantified organic residues
from each sample were strewn over four 22 x 22 mm cover slips, dried, and mounted onto microscope
slides using Norland Optical Adhesive 61. The proportion of organic residue strewn on each cover
slip (e.g. 10% of that derived from a 5 g sample) was used to calculate the equivalent mass of original
dried rock material represented (e.g. 5 g x 0.10 = 0.50 g represented on the cover slip).
To obtain an estimate of the number of palynomorphs on coverslips, the number in a 1/44 traverse of
each was counted and multiplied by 44. To give an estimate of ‗absolute abundance‘, i.e. the
concentration of palynomorphs in each sample (counts per gramme, cpg), the mean number of
palynomorphs per coverslip was divided by the approximate mass of dried rock material represented
on each coverslip. Calculations for each sample are given in Supplementary Tables B–E. Relative
abundances were estimated by counting the first three hundred palynomorphs identified (0.3% = 1
specimen; 0.7% = 2 specimens; 1% = 3 specimens, etc.). The remainder of the first coverslip and,
where applicable, the three additional coverslips, were subsequently scanned for additional rare taxa.
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Relative abundances are described as ‗rare‘ (outside the count), ‗frequent‘ (0.3–0.7%), ‗common‘ (1–
9.7%) or ‗abundant‘ (10% +). Separate counts of 100 kerogen particles (AOM, phytoclasts and
palynomorphs) were made from the un-oxidised kerogen slides. Further samples from Caistor, Bigby,
South Ferriby, Market Weighton, Bishop Wilton, East Knapton, Flixton and Speeton were processed
and analysed for palynology only, using the methods described above, with preliminary summary
results presented herein.
Standard palaeoenvironmental parameters have been calculated for the Melton Ross section,
including: (i) the ratio between terrestrial (T) and marine (M) palynomorphs (T/M ratio) as a proxy for
terrestrial input; (ii) the ratio between peridinioid and gonyaulacoid dinoflagellate cysts (P-cysts and
G-cysts, the P/G ratio) as a proxy of nutrient input; (iii) the species richness, i.e. the number of
dinoflagellate cyst taxa recorded as a proxy of their diversity. Our detailed discussion of
palaeoenvironmental parameters is provided in Eldrett et al. (2017); see also McLachlan et al. (2018)
for a recent review of the P/G ratio.
All Melton Ross palynological slides are curated in the MPA and MPK collections of the British
Geological Survey, Keyworth, Nottingham, U.K. (slide numbers MPA 70686 to 70778 and MPA
71640 to 71642; type and figured specimen numbers MPK 14662 to 14716). For the relationship
between MPA numbers and the original sample numbers used in this paper, see Supplementary Table
F. A full range chart of palynological data is available in the Supplementary Material. Full author
names and synonyms of dinoflagellate cysts, prasinophyte phycomata and acritarchs can be found in
Fensome et al. (2019).
Stable isotopes
All samples were analysed for stable carbon and oxygen isotopes on carbonates (Supplementary Table
G). Analyses were undertaken in the laboratory of the Department of Earth Sciences, Oxford
University, U.K. Oxygen and carbon isotope analytical methods were adapted from those described in
Day & Henderson (2011, section 2.7). All oxygen isotope measurements were performed on a Delta
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V Advantage isotope mass spectrometer fitted with a Gas Bench II. The Gas Bench II device
converted the carbonates to carbon dioxide (CO2) with 100% phosphoric acid (H3PO4) at 72°C
(McCrea 1950). The relative 18O/16O values (δ18O) of carbonate are expressed in per mil (‰) relative
to Vienna Pee Dee Belemnite (VPDB) on a normalised scale such that the δ18O of NBS-19 is -2.2‰
and the δ18O of NBS-18 is -23.01‰. The relative 13C/12C values (δ13C) of carbonate are expressed in
per mil relative to VPDB on a normalised scale such that the δ13C of NBS-19 is 1.95‰ and the δ13C
of NBS-18 is -5.01‰. External error (0.07 and 0.09 for δ13C and δ18O, respectively) is calculated from
repeat measurements of Oxford University‘s in-house standard NOCZ. It is assumed that the
phosphoric acid–carbonate fractionation is the same for NBS-19 and Oxford University‘s calcite
samples (Coplen 1996). For carbonates and waters, results are expressed on the same normalised scale
such that δ18O of SLAP2 reference water is -55.5‰.
Results
Organic geochemistry
The two chalk samples (top Ferriby Chalk Formation, sample -23, 0.04% TOC; Central Limestone,
sample -14, 0.10% TOC) contain the lowest organic carbon values in the sampled section. The
‗anomalous‘ lower succession marls (local Beds I–VII; samples -22 to -15) range from 0.11% to
0.16% TOC, with the exception of the darker coloured marl layer (local Bed II, sample -21), which
has 0.28% TOC. In the ‗standard‘ upper succession, the lighter coloured lithologies of Bed B (sample
-13, 0.12% TOC; sample -12, 0.20% TOC), Bed D (sample -9, 0.36% TOC) and Bed F (samples -3 to
-5, 0.12% to 0.34% TOC) contain the lowest values while the dark grey marl samples contain the
highest values: Bed C (samples -11.5, -11 and -10, 1.43% to 2.2% TOC), Bed E (samples -8.5 and -8,
2.74% and 2.18% TOC respectively) and Bed G (sample -2, 3.47% TOC). The hydrogen index (HI) in
the dark grey marl samples ranges from 78 (sample -11) to 203 mg HC/g TOC (sample -2) while the
oxygen index (OI) ranges from 15 (sample -11.5) to 26 mg CO2/g TOC (sample -8) in the same
samples (Supplementary Table H). On a modified Van Krevelen diagram (Fig. 9), these samples plot
within the Type III organic matter (mainly terrestrially derived) field, with samples -11.5 and -2
plotting towards Type III-II organic matter (mixed terrigenous and marine type). The lighter coloured
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lithologies between the dark grey marls give HI values of 40 to 78 mg HC/g TOC and relatively high
OI values (27 to 165 mg CO2/g TOC), as does sample -22 from local Bed II (HI = 39, OI = 53;
Supplementary Table H). However, with regard to characterising organic matter type, the OI data may
be unreliable in these low TOC, carbonate-rich samples, due to probable elevation by occluded CO2
within carbonate of inorganic origin, in addition to that derived from early diagenesis of organic
matter (P. Barnard pers. comm. 2019).
Palynology
In the lower succession, local Bed I (sample -22) and Bed ?III (sample -20) respectively yielded an
estimated 53 and 95 palynomorph counts per gramme (cpg) while Bed II (sample -21) has a much
higher concentration of palynomorphs, 12,408 cpg (Table 2). Beds ?IV to VII (samples -19 to -17)
yielded between 5 and 88 cpg, dominated by one taxon, Dinoflagellate? type D of Ioannides (1986)
(Table 3; Figs 6.13, 6.14). The topmost part of the lower succession (samples -15 and -16) and
overlying Central Limestone (sample -14) are palynologically barren or contain up to two
dinoflagellate cysts.
Palynological recovery and preservation from the upper succession in the freshly excavated Site 4
samples is better overall than that from many samples in other quarries investigated to date
(Supplementary Table I). Recovery from Bed B is sparse in sample -13 (28 cpg) and slightly richer in
sample -12 (288 cpg). The dark grey marl samples of the Black Band (Beds C to E) and Bed G
contain exceptionally high concentrations of palynomorphs, 85,184 to 219,648 cpg in samples -11.5, -
11, -10, 8.5, -8 and -2, i.e. in the same samples that have the highest TOC values of 1.43–3.47%. The
lighter coloured lithologies between the dark grey marls show a broad correlation between
palynomorph concentration and TOC, 17,204 to 53,284 cpg within the Black Band (samples -10.5, -9,
-7, -6), where concentration is still exceptionally high, and 1,470 to 8,668 cpg in Bed F. The
assemblage in Bed H is comparable to that in Bed G and is interpreted to be mainly derived from
reworked clasts of the latter (see Supplementary Appendix), given that Bed H is palynologically
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barren at four other localities where it has been sampled (Louth, Bigby, Caistor and South Ferriby;
Hart et al. 1993; Supplementary Table I).
The dinoflagellate cyst assemblage from local Bed II (sample -21) is diversified with 77 taxa present
(Table 2). The adjacent samples (-22 and -20) from local Beds I and ?III, along with the samples from
Bed B (-12 and -13) contain fewer taxa (32–33) but this may mainly reflect a much smaller number of
specimens inspected from these lower recovery samples. Relatively high diversity dinoflagellate cyst
assemblages are noted from the darker lithologies of the Black Band (68–79 taxa) with slightly fewer
taxa in the lighter coloured inter-beds (Bed C, sample -10.5, and Bed D; 65 and 63 taxa respectively)
and overlying Bed F (53–69 taxa). Bed G yielded 62 taxa, fewer than from comparable dark
lithologies in the Black Band.
The lower succession has the highest P/G ratio in the sampled section (average 0.61), with P-cyst taxa
Palaeohystrichophora infusorioides (Fig. 5.13) and Subtilisphaera pontis-mariae (Figs 5.14, 5.15)
respectively comprising an average 19.4% and 23.0% of total palynological assemblages from
samples -22 to -20 (Table 4). In the upper succession, P-cysts are subordinate to G-cysts in all
samples, though they occur in higher relative and absolute abundances in the more organic-rich (> 1%
TOC), dark grey marl samples (average P/G ratio of 0.33) than the lighter coloured lithologies of
Beds B–D (average P/G ratio, 0.04) and upper Bed E to Bed F (average P/G ratio, 0.26; Table 2).
Trithyrodinium suspectum (Fig. 5.12) and Ginginodinium? sp. A of Prauss (2006, 2012a) (Figs 6.3,
6.7) occur in most of the productive samples from both lower and upper successions, respectively
comprising 0.3–7% and 0.3–1.7% of total palynological assemblages. Palaeohystrichophora
infusorioides and S. pontis-mariae also occur consistently in the upper succession. With the exception
of sample -6, the former is relatively rare (1–5%) in the Black Band and more common (7.7–11.3%)
in Bed F while the latter is more common in the > 1% TOC samples of the Black Band (10.7–18.7%).
Bosedinia cf. sp. 1 of Prauss (2012b) (Figs 7.5, 7.9, 7.10) is also relatively common in the > 1% TOC
samples (1.3–6%). Eurydinium saxoniense (Fig. 5.6) occurs consistently in Beds C and D, with its
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first confirmed occurrence in sample -11.5 at the base of the former, and is relatively common (1.7–
8%) in Beds E, F and G (Table 4).
The G-cyst taxon Spiniferites spp. is prominent in most productive samples throughout the section,
5.4–28.3% of total palynological assemblages, while Pterodinium spp. comprises 1–5% of
assemblages, with an acme (10%) in sample -3 from Bed F (Table 3). Litosphaeridium siphoniphorum
(Figs 5.1, 8.12) is restricted to the lower succession, where it is common (1–2.7%). Cyclonephelium
compactum–membraniphorum (Figs 5.5, 6.4) is rare in the lower succession but is consistently
abundant throughout the upper succession, from basal Bed B to Bed H (average 21.5%). An isolated
specimen of Oligosphaeridium totum (Figs 5.9, 5.10, 8.9) was recorded from sample -17 but the taxon
is only consistently recorded from Bed B to Bed F and has an acme in upper Bed B (40.3%; Table 3).
Sepispinula? huguoniotii (Fig. 5.16) is common to abundant from Bed B to lower Bed E (samples -13
to -8.5).
There is a relative increase in prasinophyte phycomata, mainly the genera Leiosphaeridia,
Pterospermella and Tasmanites, in the darker intervals at Melton Ross (0.7–1% in Bed C; 1.3–6% in
Bed E; 6.3% in Bed G). Acanthomorph acritarchs are also present in higher percentages in the darker
> 1% TOC lithologies (average 3.9% of the total palynological assemblage) than in the < 1% TOC
samples (average 0.6%). They are mainly represented by Veryhachium spp., apart from Bed G
(sample -2), in which Micrhystridium is prominent (11%; Table 2).
Terrigenous palynomorphs are present in higher relative and absolute abundances in the more
organic-rich (> 1% TOC) samples (average T/M ratio of 0.103 and concentration of 12,555 cpg) than
in adjacent lithologies with lower (< 1%) TOC values (Table 2; average T/M ratio, 0.023; average
concentration, 524 cpg; Supplementary Table E). Terrigenous palynomorphs are mainly composed of
gymnosperm taxa (4.7–13.7% of total palynological assemblages from > 1% TOC samples),
predominantly bisaccate pollen, including Rugubivesiculites rugosus (Fig. 5.17) and Alisporites
microsaccus (Fig. 5.18), along with subordinate Classopollis spp. (Fig. 5.19) and Inaperturopollenites
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hiatus. Pteridophyte spores, including Deltoidospora spp. and Gleicheniidites spp. are persistent and
rare to common but do not exceed 3% of the total palynological assemblages. Normapolles
angiosperm pollen, Atlantopollis microreticulatus (Fig. 5.20) and Complexiopollis spp. (Fig. 5.21)
occur in most productive samples but are rare and compose less than 1% of the palynological
assemblages. With the probable exception of an occurrence of the dinoflagellate cyst Sindridinium
borealis in sample -11 (reported Albian–Early Cenomanian stratigraphic range; Nøhr-Hansen et al.
2018), palynomorphs reworked from older formations were not recorded in the Melton Ross section.
Stable isotopes
The sample (-23) from the top of the Ferriby Chalk Formation yielded a δ13C value of 2.49‰. In the
lower succession, there are three ‗peaks‘ in the δ13C data above 3‰ (samples -22 to -21, -19, and -17
to -15), separated by two ‗troughs‘ below 3‰ (samples -20, -18 to -18.5). Maximum values for the
section are from the Central Limestone (Bed A, sample -14, 4.25‰) and Bed B (sample -13, 4.21‰;
sample -12, 4.17‰). Values show an overall decline through the Black Band, from 3.91‰ (sample -
11.5) to 3.52‰ (sample -6). Above a trough in Bed F (3.34‰ in sample -4), there is a subsequent
peak in Bed G (3.77‰ in sample -2). The highest sample analysed (-1) from Bed H yielded a value of
3.14‰ (Supplementary Table G).
The δ18O data fluctuate in the -3.58 to -5.12‰ range in the lower succession. Minimum negative
values for the section occur in the Central Limestone (-3.03‰ in sample -14; Fig. 4) and overlying
basal Bed B (-3.25‰ in sample -13). Upper Bed B and Beds C to H, all yielded δ18O values more
negative than -4.5‰, with three notable peaks in the samples with > 1% TOC (lower Bed C, lower
Bed E and Bed G; -5.35‰ to -6.46‰).
Stratigraphy
Micropalaeontology
A study of foraminifera at Melton Ross has not yet been undertaken. The Cenomanian planktonic
marker taxon Rotalipora cushmani (Morrow) has been recorded from the Ferriby Chalk Formation in
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eastern England, at South Ferriby and Elsham, but has to date not been recorded from above the
erosion surface at the base of the Welton Chalk Formation (Hart et al. 1993). In most southern
England sections, it ranges up into the lower part of the Plenus Marls, e.g. Bed 3 at Dover (Jarvis et
al. 1988a) and Bed 4 at Eastbourne (Jarvis et al. 2006). At Misburg, R. cushmani occurs in
Cenomanian strata and has its last occurrence (LO) in the pre-Plenus ‗black shale‘ succession
(Hilbrecht 1986). Future inspection of our samples from Melton Ross will test for its presence in
localised Beds I–VII.
The regionally correlative Beds B–H in eastern England are dominated by a Hedbergella/Whiteinella
assemblage, with an increase in small buliminids and simple agglutinated foraminifera in the dark
coloured mudstones (Hart & Bigg 1981; Hart et al. 1993; Dodsworth 1996, fig. 6). The assemblage in
the mudstones adjacent to the Black Band is comparable to those of the upper part of the Plenus Marls
in southern England (Beds 4–8). This led Hart & Bigg (1981) to suggest that the Black Band may be
the lateral equivalent of Bed 6, a relatively clay-rich unit. On grounds of event stratigraphy, i.e. the
most argillaceous levels within the CTB successions of the Northern and Southern Provinces, Jeans et
al. (1991) also suggested correlation of eastern England Black Band, Beds C and E, with southern
England Beds 4 and 6, respectively. However, a correlation of the Black Band with a level in the
Plenus Marls is not supported by the palynological and stable isotopes data presented in this study
(see below).
Hart & Bigg (1981) found the first occurrence (FO) of Helvetoglobotruncana praehelvetica (Trujillo)
in Bed H at Elsham. This taxon is an early morphotype of Helvetoglobotruncana helvetica (Bolli), the
diagnostic zonal marker for the Early to Middle Turonian in Tethyan areas. The FO of H. helvetica
was tentatively recorded from a marl seam at South Ferriby (Hart & Leary 1989) that probably
correlates with the marl seam above the Turnus Bed at Melton Ross (Fig. 4). However, its FO is based
on the (highly subjective) evolutionary boundary between praehelvetica and helvetica (Hart & Leary
1989). The rarity of H. helvetica at high latitudes such as Lincolnshire also renders its FO
impracticable as a confident time-diagnostic event in the Northern Province (Hart 2019). In NW
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Germany, Hilbrecht (1986) reported the FO of H. helvetica within the Fish Shale. At Dover, southern
England, its FO is difficult to locate precisely in the nodular chalk lithology of the Ballard Cliff
Member (above the Plenus Marls), as this is difficult to process for calcareous microfossils (Hart &
Leary 1989). At Eastbourne, the FO of H. helvetica is picked between Mead Marls 3 and 4 of the
Ballard Cliff Member (Jarvis et al. 2006). In Fig. 4, the planktonic foraminiferal zonation is
extrapolated to Melton Ross from Elsham and South Ferriby.
Palynology
Several dinoflagellate cysts have widespread range bases and tops in the Cenomanian and Turonian
stages, and have been used to zone their substages (Clarke & Verdier 1967; Burgess 1971; Foucher
1981; Williams 1977; Williams et al. 2004). Regional zonations have recently been revised for
Central and Northern Europe (Olde et al. 2015a) and adapted for the Western Interior of the USA
(Dodsworth & Eldrett 2019).
The LO of consistent and common Litosphaeridium siphoniphorum in the lower succession at Melton
Ross (sample -20) correlates with its LO in the pre-Plenus sequence of NW Germany, at Wunstorf
outcrop (Marshall & Batten 1988) and core (van Helmond et al. 2015), and Misburg outcrop
(Marshall & Batten 1988). In southern England, the LO of common L. siphoniphorum occurs in
Plenus Marls Bed 6 at Lulworth and Eastbourne (Fig. 1; Dodsworth 2000; Pearce et al. 2009) while
isolated specimens have been recorded from Bed 7 and Bed 8 at Lulworth (Dodsworth 2000). Isolated
specimens have also been sporadically recorded from Turonian and Coniacian deposits at some
localities in southern England, France, NW Germany (Clarke & Verdier 1967; Foucher 1982, 1983;
Marshall & Batten 1988) and Pueblo, Colorado (Dodsworth 2000). These isolated Turonian and
younger occurrences are probably reworked specimens. Previous reports of the LO of consistent L.
siphoniphorum in Lower Turonian deposits (Williams & Bujak 1985; Costa & Davey 1992) probably
derive from former assignment of the Plenus Marls in southern England to this substage (e.g. Jefferies
1962, 1963); the Plenus Marls have subsequently been confidently reassigned to a Late Cenomanian
age (e.g. Jarvis et al. 1988a; Gale et al. 1993). A Late Cenomanian LO of consistent L. siphoniphorum
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has been reported worldwide, including other locations in southern England (Davey 1969; Hart et al.
1987; Jarvis et al. 1988b), the Witch Ground Graben, central North Sea (Harker et al. 1987), France
(Foucher 1979, 1980; Courtinat et al. 1991), northern Spain (Mao & Lamolda 1999), Crimea
(Dodsworth 2004a), Poland (Dodsworth 2004b), eastern USA (Aurisano 1989), the Western Interior
of the USA (Courtinat 1993; Dodsworth 2000, 2016; Eldrett et al. 2015a; Dodsworth & Eldrett 2019),
Ocean Drilling Project holes including Demerara Rise (Fig. 1; Leg 207, Site 1260) and Kerguelen
Plateau (Leg 183, Site 1138; Eldrett et al. 2017), Australia (Morgan 1980; McMinn 1988) and New
Zealand (Hasegawa et al. 2013; Schiøler & Crampton 2014).
At Melton Ross, the LO of Pterodinium crassimuratum (Fig. 5.7) occurs in sample -21, close to the
LO of consistent/common L. siphoniphorum in sample -20. Davey & Williams (1966) and Clarke &
Verdier (1967) recorded it from beds of Middle and Late Cenomanian age in southern England, its LO
coinciding with that of L. siphoniphorum within the Plenus Marls on the Isle of Wight and at a
slightly higher level at Eastbourne, in Plenus Marls Bed 8 (Pearce et al. 2009). At Pueblo, Colorado,
the LO of P. crassimuratum also occurs in the Upper Cenomanian (Dodsworth & Eldrett 2019).
The FO of abundant Cyclonephelium compactum–membraniphorum in lower Bed B (sample -13),
immediately above the Central Limestone at Melton Ross, correlates with the base of its abundant
occurrence immediately above the Plenus Bed in NW Germany, at Wunstorf outcrop (Marshall &
Batten 1988) and core (van Helmond et al. 2015), and Misburg outcrop (Marshall & Batten 1988).
The event is less clearly defined in southern England, but can be tentatively picked near the top of the
Plenus Marls, in Bed 7 at Eastbourne (Pearce et al. 2009) and Bed 8 at Lulworth (Dodsworth 2000).
Cyclonephelium membraniphorum is rare or absent in coeval Tethyan sections farther south, in Spain
(Lamolda & Mao 1999; Peyrot 2011; Peyrot et al. 2011, 2012) and Morocco (Prauss 2012a, b).
At Melton Ross, an influx of Oligosphaeridium totum from Bed B to lower Bed C can be correlated
with an influx of O. totum from the same levels at Caistor (samples CL-11 to CL-9; 1–3%) and from
upper Bed B at Market Weighton (samples MW-20 to MW-18; 3–6%; see Supplementary Table I for
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sample positions). Duane (in Hart et al. 1993) recorded its highest relative abundance (3.4%) in the
lower part of the Black Band (their sample SFE-18) at South Ferriby (the underlying Bed B samples
SFE-19 and SFE-20 at South Ferriby were palynologically barren). In NW Germany, Marshall &
Batten (1988) only recorded O. totum from above the Plenus Bed at Wunstorf (their sample 38) and at
Misburg (their samples 27, 22 and 20). At Misburg, an influx of O. totum was indicated, with its base
in sample 22, collected from a bed that contains the only occurrence of the latest Cenomanian zonal
ammonite Neocardioceras juddii in the section. An acme of O. totum occurs in overlying sample 20.
In central Poland, O. totum occurs in sample Pul-17 of the Pulawy borehole (Fig. 1), within the δ13C
excursion that spans the CTB (Peryt & Wyrwicka 1993), one sample above the FO of abundant C.
compactum–membraniphorum (sample Pul-16). This, in turn, is stratigraphically higher than the LO
of consistent/common L. siphoniphorum (sample Pul-15). Thus, the three bio-events also occur in the
same relative order over a condensed (< 1 m thick) interval in that section (Dodsworth 2004b).
The LO of Adnatosphaeridium tutulosum (Fig. 5.4) occurs in Bed F at Melton Ross (sample -5). It
was recorded from Bed C at Flixton (Dodsworth 1996). At other European and North America
locations, the LO of A. tutulosum also occurs within the uppermost Cenomanian, above the LO of
consistent/common L. siphoniphorum and the FO of abundant C. compactum–membraniphorum: in
NW Germany, above the Plenus Bed at Wunstorf and Misburg outcrops (Marshall & Batten 1988); in
southern England, at Hooken Cliffs, Devon (Jarvis et al. 1988b), Eastbourne (Plenus Marls Bed 8;
Pearce et al. 2009) and 0.5 m above the Plenus Marls at Lulworth (Dodsworth 2000); and in southern
France (cf. Courtinat et al. 1991 & Jarvis et al. 2011) and the Western Interior of the USA
(Dodsworth 2000; Harris & Tocher 2003; Dodsworth & Eldrett 2019). The LO of A. tutulosum is at
the same level as the LO of L. siphoniphorum in northern France (Foucher 1983) and Crimea
(Dodsworth 2004a).
The LO of Carpodinium obliquicostatum (Fig. 5.3) occurs in Bed C (sample -11.5) at Melton Ross. It
was recorded from Bed E at South Ferriby and Flixton (Dodsworth 1996). In NW Germany, Marshall
& Batten (1988) reported it above the Plenus Bed in the Wunstorf and Misburg outcrops. In southern
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England, it occurs 0.5 m above the Plenus Marls at Lulworth (Dodsworth 2000). Although rare and
sporadic in occurrence, its range top in the Western Interior is reported to coincide approximately
with that of A. tutulosum at Pueblo, Colorado (Dodsworth 2000).
Microdinium setosum (Fig. 6.6) occurs fairly consistently in the Black Band at Melton Ross, with its
LO in Bed E (sample -6). In southern England, it occurs throughout the Cenomanian (Clarke &
Verdier 1967) and is sporadic in the Plenus Marls with an LO in Bed 2 at Eastbourne (Jarvis et al.
2011) and Bed 7 at Lulworth (Dodsworth 2000). However, it occurs sporadically as high as Middle
Turonian in the North Sea Basin (Costa & Davey 1992).
At Melton Ross, Adnatosphaeridium? chonetum (Fig. 5.8) was recorded from local Bed II (sample -
21), with an isolated, questionable specimen also present in Bed E (sample -6). It has previously been
recorded from Bed C at Flixton (Dodsworth 1996). The occurrence of A.? chonetum in Cenomanian
deposits was noted by Davey (1969) in Northern Europe and by Cookson & Eisenack (1962) and
Backhouse (2006) in Australia. In the Western Interior of the USA, it is common in the Upper
Cenomanian Substage, with an uppermost Cenomanian LO (Harris & Tocher 2003; Dodsworth &
Eldrett 2019). In the Shetland Group, offshore Norway, it reappears in Coniacian and Lower
Santonian deposits (PD, personal observation).
The FOs of Canningia glomerata, Heterosphaeridium difficile, Florentinia buspina, F.? torulosa and
Senoniasphaera turonica are intra-Lower Turonian biostratigraphic marker events in Europe (Davey
& Verdier 1976; Foucher 1980, 1981, 1983; Tocher & Jarvis 1987; Jarvis et al. 1988a; Costa &
Davey 1992; FitzPatrick 1995; Pearce et al. 2003, 2009, 2011). These taxa have not been recorded at
Melton Ross or in any of the other eastern England localities/samples indicated in Supplementary
Table I. Thus, Beds C–G are probably stratigraphically lower than their range bases.
In terms of dinoflagellate cyst zonation, the palynologically productive samples from the lower
succession at Melton Ross are assigned to the Litosphaeridium siphoniphorum Interval Zone of Olde
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et al. (2015a) and Dodsworth & Eldrett (2019). The productive part of the upper succession (Bed B to
Bed G) is assigned to the Cyclonephelium membraniphorum Zone of these authors, with Bed B to
lowermost Bed F belonging to the Adnatosphaeridium tutulosum Subzone of Dodsworth & Eldrett
(2019; Fig. 4).
At Wunstorf, Prauss (2006) reported acanthomorph acritarchs. Higher relative abundances of
Veryhachium (ca. 1–2%) occur in darker lithologies from the pre-Plenus and lower post-Plenus
succession, relative to lighter coloured inter-beds. There is a large influx of Micrhystridium in the
upper part of the Fish Shale (ca. 15–70%). Following Wall (1965) and Downie et al. (1971), Prauss
(2006) attributed the Veryhachium occurrences to a distal offshore, hydrodynamically quiet
environment, and the prominent acritarch peak dominated by Micrhystridium within the upper part of
the Fish Shale to the influence of a relatively near-shore turbulent water environment. An up-section
change from common Veryhachium (Beds C to E) to abundant Micrhystridium (Bed G) at Melton
Ross may correlate with that reported from Wunstorf.
The consistent presence of the bisaccate pollen Rugubivesiculites rugosus in the upper succession at
Melton Ross is noteworthy. It has recently been reported throughout the Upper Cenomanian–Middle
Turonian section exposed at Pueblo, Colorado (Dodsworth & Eldrett 2019) and in Turonian cores
from Texas (e.g. Iona-1; Eldrett et al. 2017). In the Greenland area, Nøhr-Hansen (2012, appendix 1),
Fensome et al. (2016) and Nøhr-Hansen et al. (2016) indicated that Rugubivesiculites spp. / R.
rugosus has a consistent LO within the Turonian.
Calcareous nannofossils
A study of calcareous nannofossils at Melton Ross has not yet been undertaken, but Bralower (1988)
reported their ranges from the CTB interval at other quarries, including the nearby sections at South
Ferriby and Elsham (Fig. 2). The LOs of the coccoliths Axopodorhabdus albianus (Black), Helenea
chiastia Worsley and Rhagodiscus asper Stradner occur in Bed H at South Ferriby. At Elsham, the
LO of R. asper also occurs in Bed H but the LOs of H. chiastia and A. albianus occur respectively 20
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cm and 15 cm lower, within undifferentiated variegated marls. At Misburg, NW Germany, all three
events occur within Lower Turonian deposits above the Fish Shale (cf. Bralower 1988, fig. 21, and
Hilbrecht & Hoefs 1986, fig. 2), with the LOs of H. chiastia and A. albianus in a ‗triple band‘ of
stratigraphically higher, dark-coloured mudstones, and the LO of R. asper ca. 5 m above the ‗triple
band‘. At Dover, southern England, the three events occur stratigraphically lower, within Upper
Cenomanian deposits, i.e. the highest 0.55 m of the Plenus Marls which is 4.45 m thick at the sampled
location. At Pueblo, Colorado, the three events occur over a ca. 4.5 m interval that spans the CTB, in
the stratigraphic order of LO A. albianus, LO H. chiastia and LO R. asper (Fig. 10). Bralower (1988)
suggested that slow sedimentation rates, a probable hiatus between Bed H and the Turnus Bed, and
subsequent bioturbation may have ‗smeared‘ these events at South Ferriby and Elsham, while at
Dover they occur in proximity to a lithological change to the hard, condensed limestone of the Ballard
Cliff Member that overlies the Plenus Marls.
Stable isotopes
Before discussing the possible stratigraphic significance of the measured changes in the δ13C stable
isotope record, diagenetic or local lithological influences have to be excluded. Diagenetic alteration is
commonly observed in fine-grained carbonate sedimentary rocks. Deep burial cementation and
recrystallization can result in the addition of isotopically depleted calcite to the bulk carbonate pool,
shifting the bulk δ18O record towards lower values (e.g. Jarvis et al. 2011, 2015). However, during
burial diagenesis, carbon isotope values are less prone to diagenetic alteration than oxygen isotope
values as the carbon isotope system is rock-dominated and δ13C is subject to a much smaller
temperature-controlled fractionation (Marshall 1992). A cross-plot of δ13C and δ18O allows the
elucidation of the depositional and post-depositional controls on stable isotope values, with a strong
co-variance between δ13C and δ18O indicating potential diagenetic influence (Fig. 11).
In general, the stable isotope results for the Melton Ross section show poor co-variance (R2= 0.0076),
indicating limited diagenetic overprint. However, the samples from Beds I–VII and Beds A and B
that contain low TOC (< 0.2%) do show a relatively strong co-variance (R2= 0.84), indicating
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potential dissolution and re-precipitation of isotopically light 13C cements through interaction with
meteoric pore fluids during burial in this interval. Conversely, the δ13C and δ18O co-variance remains
poor in samples with low TOC from Beds C–H. It is uncertain why the lower part of the succession
may be preferentially affected by diagenesis. The range of δ13C and δ18O values from Melton Ross
appear consistent with bulk stable isotope values from coeval European Cenomanian–Turonian
sections that record a 2–5‰ positive carbon isotope excursion (CIE) associated with OAE-2 (Fig. 11).
The δ13C profile from Beds A–H at Melton Ross broadly correlates with that assigned to OAE-2 at
South Ferriby (Schlanger et al. 1987; Hart et al. 1991; Clarkson et al. 2018), which also shows
highest δ13C values in Beds A and B and an overall decreasing trend in values through Beds C to H,
with a relatively minor, terminal peak around Bed G/H. The South Ferriby studies sampled metres of
chalk below and above Beds A–H, and indicated ca. 1‰ lower δ13C background values below and ca.
0.5‰ lower background values above. Therefore, although the δ18O record may have been influenced
by diagenesis, the δ13C signal appears less impacted and possibly reflects a primary
palaeoceanographic signal. A similar conclusion was drawn by Hu et al. (2012) and Mitchell (2019)
for the chalks and marls from the Northern Province.
The detailed profile of the CIE associated with OAE-2 has been shown to correlate across the
Southern Province in England, including Dover and Eastbourne (Jarvis et al. 2006), with Eastbourne
being taken as the principal reference section (Paul et al. 1999). It has also been correlated inter-
regionally, particularly with the USA (e.g. Gale et al. 1993; Joo & Sageman 2014; Eldrett et al.
2015a; Fig. 10), Morocco (Tsikos et al. 2004; Jenkyns et al. 2017), Japan (Uramoto et al. 2013) and
other areas of Europe (Jarvis et al. 2015). In Misburg outcrop (Hilbrecht 1986), Gröbern core (Voigt
et al. 2006) and Wunstorf core (van Helmond et al. (2015), NW Germany, background δ13C values
are recorded in the pre-Plenus sequence while peak values associated with OAE-2 occur within the
Plenus Bed. The new record of peak δ13C values in the Central Limestone at Melton Ross supports its
correlation with the Plenus Bed, as initially predicted by Wood & Mortimore (1995). The magnitude
and top of the δ13C excursion vary across NW Germany (Hilbrecht et al. 1992), but it shows an
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overall decreasing trend through the Fish Shale at Misburg and Wunstorf, possibly correlating with
that seen through Beds C to H at Melton Ross and South Ferriby.
In the Southern Province of England, a ‗build-up phase‘ of overall increasing δ13C values through
Beds 1–2 of the Plenus Marls precedes a lower peak (‗a‘) around Bed 3, a relatively thick limestone
unit. Praeactinocamax plenus occurs in the overlying part of the Plenus Marls (Beds 4–8),
particularly in Bed 4 (Jefferies 1962, 1963). Above a trough in δ13C values through Beds 4–8, a
second and maximum peak (‗b‘) occurs close to the boundary between the Plenus Marls and overlying
Ballard Cliff Member. Above peak ‗b‘, δ13C values remain high but there is an overall decline in
values across the CTB, through the Ballard Cliff Member, including some prominent marl seams
(Mead Marls 1–6), below a final OAE-2 peak ‗c‘ (Jarvis et al. 2006).
In terms of correlation between the Northern Province and Southern Province in England, our new
δ13C data from Melton Ross could be interpreted as follows. The three (> 3‰) peaks of δ13C in Beds
I–VII may be ‗precursor‘ / build-up events to OAE-2, correlating with southern England Beds 1–2;
the base of the > 4‰ interval in Bed A (Central Limestone) equals peak ‗a‘ of Jarvis et al. (2006;
southern England Bed 3); lower Bed B may correlate with part of Beds 4–8 in southern England,
supported by the record of P. plenus at Melton Ross, Site 1 (Wood et al. 1997; Fig. 4); the top of the >
4‰ interval in upper Bed B equals peak ‗b‘ of Jarvis et al. (2006); the trough in Beds C–F and peak
in Bed G may correlate with the trough through the Ballard Cliff Member and peak ‗c‘ in southern
England (Fig. 10). A comparable correlation has been made between the δ13C profile from Beds A–H
at South Ferriby and that of the Southern Province (Wood & Mortimore 1995; Clarkson et al. 2018).
Notwithstanding concerns about diagenetic alteration of the eastern England Chalk Group oxygen
isotope signal discussed above, the δ18O curve from Melton Ross (Fig. 4) is comparable to those from
other parts of Europe (cf. Jarvis et al. 2011, fig. 8). The minimum values in the Central Limestone and
overlying lower Bed B may correlate with the PCE. The higher values from Beds C–H could be
considered consistent with subsequent warming around the CTB. The peaks in δ18O values from the
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most organic-rich layers could reflect temperature maxima and/or increased run-off during their
deposition (see below).
Discussion
Organic geochemistry
In an overview of OAE-2 in North European shelf settings, Jenkyns (1985) reported TOC values
mainly within the range of 1–3% and HI from 150–300 mg HC/g TOC (mixed marine and terrigenous
sources) in Cenomanian–Turonian organic-rich deposits, including eastern England (South Ferriby,
0.79–3.15% TOC) and NW Germany (1.2–2.8% TOC; Schlanger et al. 1987). The new HI data from
the > 1% TOC dark mudstones at Melton Ross plot around the lower end of this range (78–203 mg
HC/g TOC; Fig. 9), consistent with a relatively large terrigenous component. Jenkyns (1985) noted
that there appears to be a general trend of an increase in preservation of organic carbon as the
depositional environment deepened from stable, relatively shallow shelf to rifted graben and deeper
continental margin. Higher TOC values have since been reported from the Black Band farther north in
eastern England, from Speeton (13% TOC, Farrimond et al. 1990) and Flixton (4.5–10.2% TOC,
Jeans et al. 1991) at the edge of the Cleveland Basin (Fig. 2). Herbin et al. (1986) reported 30% TOC
from the Black Band Bed in the Central Graben of the North Sea, including large quantities of Type
III organic matter, consistent with a substantial terrigenous component. In palynological assemblages
from Melton Ross and other onshore Black Band localities, marine organic-walled phytoplankton
dominate over terrigenous pollen and spores (Supplementary Table I), although at Flixton, the latter
compose approximately one half of the assemblages from Bed C.
The Cyclonephelium compactum–membraniphorum issue
The southward incursion of abundant Cyclonephelium compactum–membraniphorum into Central
Europe has been attributed to a stressed marine environment associated with oceanic anoxia (Marshall
& Batten 1988; Courtinat et al. 1991; Hart et al. 1991). However, its initial migration has more
recently been correlated with the PCE, including sites outside Europe. The latter include a proto-
Atlantic coastal setting in New Jersey (Bass River; van Helmond et al. 2014; Fig. 1) and the Western
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Interior Seaway of the USA (e.g. Portland-1 core, central Colorado; Iona-1 core, SW Texas; Eldrett et
al. 2014, 2017; van Helmond et al. 2016; Fig. 1). At Melton Ross, its basal abundant occurrence in
sample -13 is at the same level as records of Praeactinocamax plenus (Fig. 4) and it is consistently
abundant up to the top of the sampled section (sample -1). While the initial southward migration may
have been linked to the PCE, its subsequent abundance in latest Cenomanian and Early Turonian
deposits occurred during a time of extreme warmth. Its post-PCE prosperity during the OAE-2
interval may be related to the relative tolerance of areoligeracean dinoflagellate cysts in general to
stressed marine environments, where their abundant occurrence is often associated with nearshore
environments / falls in relative sea level (Brinkhuis & Zachariasse 1988; Harker et al. 1990, p. 202-
204; Li & Habib 1996; Olde et al. 2015b). In this case, the environmental stress is possibly associated
with oxygen-deficient conditions in the water column and photic zone (Marshall & Batten 1988)
and/or increased runoff (van Helmond et al. 2015).
Samples with prominent Dinoflagellate? type D of Ioannides (1986)
Dinoflagellate? type D occurs consistently throughout the sampled section at Melton Ross, and in
dominant relative abundance (48–100% of palynological assemblages) in the poorly preserved
material from samples -20 to -17, from the local Beds III?–VII. However, inspection of the absolute
abundance data (Table 3) indicates that its occurrences in these beds are in the single units to tens per
gramme range, less than its hundreds per gramme concentration estimates from beds with richer
recovery. It is suggested here that its dominance in Beds III?–VII is due to its preferential
preservation, probably related to a relatively thick and resilient wall structure. This interpretation is
supported by the data of Batten (in Wood et al. 1997), who reported diversified dinoflagellate cyst
assemblages with L. siphoniphorum up to local Bed VI at Melton Ross, Site 2, i.e. in correlative strata
a few tens of metres away (Fig. 3).
Weathering of exposed quarry sections probably destroys palynomorphs and other organic matter
over a short time, of months to a few years, with only central levels within the Black Band being
unaffected at some other sampled localities, e.g. South Ferriby and Caistor. Weathered samples
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contain lower concentrations of poorly preserved palynomorphs, mainly those with relatively thick
walls: Dinoflagellate? type D, Kalyptea spp., prasinophyte phycomata and indeterminate peridinioid
endocysts. Samples worst affected are marked with an asterisk in Supplementary Table I.
Terrigenous input
In eastern England, the higher T/M ratio in relatively organic-rich (> 1% TOC) lithologies
documented here from Melton Ross (Table 2) has previously been noted in Black Band samples from
Flixton (Dodsworth 1996, fig. 9; T/M ratio 0.27–0.54 in 4.5–10.2% TOC samples, and 0–0.04 in 0.1–
0.4% TOC samples). Analyses from other Yorkshire sections (Fig. 2) appear to confirm this pattern:
East Knapton (T/M ratio average 0.11 in dark lithologies and 0.02 in light lithologies), Market
Weighton (T/M ratio average 0.08 in dark lithologies and 0.01 in light lithologies) and, to a lesser
extent, Bishop Wilton (T/M ratio average 0.06 in dark lithologies and 0.04 in light lithologies;
Supplementary Table I). This trend cannot be discerned at other Lincolnshire localities (South
Ferriby, Bigby and Caistor), where Bed B and Bed F/G samples are heavily weathered or barren and
Bed D is relatively argillaceous and tentatively picked on slightly less dark marls within the Black
Band.
At Wunstorf, Prauss (2006) calculated an average T/M ratio of ca. 0.2 for the CTB succession. Prauss
(2006) and van Helmond et al. (2015) also reported elevated relative and absolute abundances of
spores and pollen in dark coloured mudstones compared with interbedded organic-poor lithologies,
particularly within the Fish Shale (T/M ratio ca. 0.2–0.46, in ca. 1–3% TOC rocks). Van Helmond et
al. (2015) interpreted this pattern as indicative of an intensified hydrological cycle during OAE-2, i.e.
increased evaporation, precipitation and run-off during ‗black shale‘ deposition, associated with
global warming after the PCE (see also Heimhofer et al. 2018). Reworking of mud to the basin (e.g.
Jenkyns 1980) and/or the formation of swamps due to drowning of land masses (cf. Ioannides et al.
1976) during phases of CTB transgression could also have contributed to the increase in spores and
pollen (Dodsworth 2000).
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In eastern England, gymnosperm bisaccate pollen are invariably the dominant terrigenous
palynomorph group in the Black Band and Bed G (Dodsworth 1996; this paper). The productive
samples from the lower succession at Melton Ross (samples -22 to -20; Table 2) contain few
terrigenous palynomorphs (T/M ratio 0.01 or less) which are also mainly bisaccate pollen. At
Wunstorf, however, saccate pollen constitute less than one half of the terrigenous palynomorphs in
most samples from the CTB succession, with trilete spores and non-saccate pollen often composing
most of the assemblage (Prauss 2006; van Helmond et al. 2015). This contrast between the regions
probably reflects greater proximity to sources of detrital input in the Lower Saxony Basin, as reflected
by a much thicker succession there. Prauss (1993) noted that trilete spores show higher relative
abundances than saccate pollen in marine deposits with greater fluvial input because the latter are
more readily transported by wind (e.g. Mudie & McCarthy 1994). Different hydrodynamic properties,
i.e. the relative buoyancy of saccate pollen, is probably also a factor in their differential sorting with
distance from fluvial-deltaic sources (cf. Muller 1959). In the Cenomanian deposits of Texas, spores
form much larger proportions of terrigenous palynomorph assemblages in the proximal East Texas
Basin than in the relatively distal Maverick Basin (Dodsworth 2016).
Pollen provincialism
Herngreen & Chlonova (1981), Herngreen et al. (1996) and Traverse (2007) described Late
Cretaceous palynofloral provinces. Costa & Davey (1992) reported the southerly limit of triprojectate
pollen (the Aquilapollenites Province) to correlate approximately with the northerly siliciclastic
Shetland Group – southerly calcareous Chalk Group transition in Northern Europe, around modern
latitude 59°N. The rare but consistent occurrences of Normapolles pollen from the CTB succession at
Melton Ross complement previous records of their presence in the Black Band at Flixton, South
Ferriby and North Sea well 47/10-1 (Fig. 2; Dodsworth 1996). These may be the most northerly
published Cenomanian–Turonian records in Europe (cf. Peyrot et al. 2008, fig. 4) and indicate
assignment of the eastern England Chalk Group to the Normapolles Province. Normapolles pollen are
a more common component of palynological assemblages from coeval deposits farther south in
Europe (e.g. Bulgaria, Pavlishina & Minev 1998; Romania, Ion et al. 2004; SE France, Heimhofer et
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al. 2018; Spain, Peyrot et al. 2008, 2011). In SE France, Heimhofer et al. (2018) suggested that the
PCE may have fostered a first spread of Normapolles-type angiosperms. Climatic conditions would
have been cooler and less humid during the PCE, possibly resulting in an open, savanna-type
vegetation community with increased abundances of Normapolles-producing angiosperms.
Phytoplankton productivity and preservation
The P-cyst Palaeohystrichophora infusorioides and the G-cyst Spiniferites ramosus constitute the
main components of European dinoflagellate cyst assemblages from Late Cretaceous offshore/deeper
water environments and upwelling zones (Pearce et al. 2003; Olde et al. 2015b). The P-cyst genera
Palaeohystrichophora and Subtilisphaera have been suggested as heterotrophic taxa. High numbers
may reflect increased nutrient availability from upwelling in the Atlantic Ocean off the NW African
(Prauss 2012b) and European margins, and subsequent transport of nutrients via a proto-Gulf Stream
to the Anglo-Paris Basin (Prince et al. 1999, 2008; Pearce et al. 2009). Subtilisphaera pontis-mariae
and P. infusorioides mainly compose the high P/G ratio in Melton Ross local Beds I–?III (average
0.61). A source for enhanced nutrient supply during this interval is uncertain. The low T/M ratio is
inconsistent with increased run-off. Upwelling as a nutrient source requires a thermal difference and
sea floor topographical variation but there is no evidence for these in eastern England.
Phytoplankton productivity during OAE-2 ‗black shale‘ deposition has previously been discussed
from Wunstorf. Linnert et al. (2010) studied assemblages of calcareous nannofossils, showing a shift
from a generally oligotrophic ecosystem during deposition of lighter coloured beds to more
mesotrophic or even eutrophic conditions during deposition of dark coloured mudstones. Van
Helmond et al. (2014, 2015) suggested that an acceleration of the hydrological cycle during the
warmer intervals of OAE-2 may have played a key role in supplying nutrients offshore and enhancing
stratification, thus contributing to the development of ocean anoxia. Increased precipitation, run-off
(reflected by a higher T/M ratio) and associated nutrients may have contributed to productivity of
organic-walled phytoplankton during deposition of the dark coloured layers in eastern England.
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Within the Black Band and Bed G at Melton Ross, common Eurydinium saxoniense and Bosedinia cf.
sp. 1 of Prauss (2012b) and rare Bosedinia laevigata, make a greater contribution to the P/G ratio in
the > 1% TOC samples (Table 4). Eurydinium saxoniense was initially described in higher relative
abundances in dark coloured mudstone layers from Wunstorf and Misburg by Marshall & Batten
(1988), who associated it with a stressed marine environment with a low level of oxygen extending
high up the water column.
High relative abundances of Bosedinia spp. in Late Cretaceous marine settings have been interpreted
as reflecting reduced salinity surface water and enhanced density stratification (at Tarfaya; Prauss
2012b, fig. 5), and/or increased nutrient (nitrite/nitrate) availability from oxygen-deficient waters
encroaching into the photic zone (in the Western Interior of the USA and Demerara Rise; Dodsworth
2016; Eldrett et al. 2017). In the Black Band and Bed G, higher numbers of E. saxoniense and
Bosedinia spp. occur in the absence of fresh/brackish-water algae and may record some stimulation of
dinoflagellate productivity by increased supply of reduced nitrogen chemo-species in photic-zone
waters (cf. Prauss 2007). This is supported by the relative increase in prasinophyte phycomata in the
darker layers at Melton Ross, Flixton (5.7–9.3% in Bed C, 14% in Bed E; Dodsworth 1996) and
Wunstorf (up to 15%; Prauss 2006). Prasinophyte prosperity may be mainly related to productivity
stimulated by introduction of nitrogen/ammonium-enriched waters of the denitrification zone into the
photic zone (Prauss 2006, 2007). However, the average P/G ratio at Melton Ross (0.33) in the > 1%
TOC samples is much lower than that discussed above for Bosedinia-dominated organic-rich
lithologies from, for example, the Cenomanian of the Western Interior (ca. 0.75–0.85), and may
therefore, reflect a relatively moderate increase in peridinioid productivity. Likewise, prasinophyte
increases are moderate relative to uppermost Cenomanian organic-rich mudstones in Aksudere,
Crimea (Fig. 1; up to 7.2% TOC; Naidin 1993), where they typically constitute over half the
palynomorph assemblage (Dodsworth 2004a). Regardless of the relative contribution of organic-
walled phytoplankton productivity to OAE-2, the higher proportions of E. saxoniense, Bosedinia and
prasinophytes in the > 1% TOC lithologies at Melton Ross and Flixton are consistent with increased
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quality of sea floor preservation in a low O2 environment. They may indicate periodic
hydrographically restricted and eutrophic conditions (cf. Eldrett et al. 2017).
Palynomorph concentration
Most palynological studies of Cenomanian and Turonian deposits report relative abundances only.
Records of specimen numbers per microscope slide from Dover, Eastbourne and the Isle of Wight,
southern England (Jarvis et al. 1988a; Fitzpatrick 1996), have given a semi-quantitative indication of
recovery. Until the last ten years, there have been few reports of fully quantitative concentration,
counts per gramme (cpg) data. Duane (in Paul et al. 1994), using a volumetric method, reported
dinoflagellate cysts in the hundreds (mainly 100–800 cpg) range from Middle Cenomanian chalks and
marls in the Dover–Folkestone Warren area, southern England. Pearce et al. (2003), using a method
involving adding a known number of Lycopodium spores to samples (Stockmarr 1971; Mertens et al.
2009), reported dinoflagellate cysts in the tens to 600 cpg range from Turonian chalks and marls at
Banterwick Barn, southern England.
Much higher concentrations of palynomorphs (mainly dinoflagellate cysts) were reported from Upper
Cenomanian deposits in southern England, at Eastbourne and Lulworth. Pearce et al. (2009), using the
Lycopodium method, reported ca. 50,000–75,000 cpg from six marl and chalk samples over a 10 m
interval below the Plenus Marls at Eastbourne. Dodsworth (2000), using this study‘s volumetric
method, reported nearly 38,000 cpg from a marl sample one metre below the Plenus Marls at
Lulworth. Pearce et al. (2009) noted that assemblages from below the Plenus Marls at both localities
are dominated by the P-cyst Palaeohystrichophora infusorioides. They attributed its high
concentrations to upwelling in the Atlantic Ocean off the European and NW African margins, with
associated nutrients being transported to the southern England area by dominantly south-westerly
surface winds and ocean surface currents. Both Dodsworth (2000) and Pearce et al. (2009)
documented overall decreasing concentrations of palynomorphs through the Plenus Marls (to just over
100 cpg in Bed 8 at Lulworth) and barren or impoverished recovery in the overlying Ballard Cliff
Member. Jarvis et al. (1988a), FitzPatrick (1996) and Pearce et al. (2003) also reported a
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palynologically barren or impoverished interval above the Plenus Marls at other southern England
localities, with better palynological recovery from Turonian chalks and marls above the Ballard Cliff
Member and Holywell Member.
Dodsworth (2000) attributed poor palynological recovery from the upper part of the Plenus Marls and
the Ballard Cliff Member to upward coarsening through the succession (Hancock 1989; Jeans et al.
1991; Lamolda et al. 1994) and the associated reduced palynomorph preservation potential of
lithologies with a relatively high coarse silt and sand grade component. Pearce et al. (2009)
acknowledged this but, following Jarvis et al. (1988a), FitzPatrick (1996) and Lamolda & Mao
(1999), suggested that a primary decline in phytoplankton productivity across the CTB in the Anglo-
Paris Basin may have been an additional cause of the low recovery. The Eastbourne and Lulworth
localities record a marked decrease in the relative and absolute abundance of P. infusorioides above
Bed 4 of the Plenus Marls, consistent with a decrease in nutrient supply (Pearce et al. 2009). An
overall decline in the nannofossil fertility index also occurs between Bed 4 and Ballard Cliff Member
(Gale et al. 2000).
Palynomorph concentration data are becoming more frequently available from areas outside southern
England. From the CTB succession at Bass River, New Jersey, van Helmond et al. (2014), using the
Lycopodium method, reported dinoflagellate cysts in the 2,000–30,000 cpg range, and terrigenous
spores and pollen in the 400–11,000 cpg range, from silty clay to clayey silt lithologies (0.61–1.51%
TOC) that were deposited in an inner neritic setting. From the CTB succession in Wunstorf core, van
Helmond et al. (2015), also using the Lycopodium method, reported dinoflagellate cysts mainly in the
1,000–15,000 cpg range and terrigenous spores and pollen in the 2,500–5,000 cpg range, from
samples with 0.5–3% TOC. Using this study‘s volumetric method, comparable values for organic-
walled phytoplankton were obtained from the CTB succession at Aksudere, Crimea, 3,520–20,400
cpg, from samples with 0.3–7.2% TOC (Dodsworth 2004a). Comparable values have also been
obtained from relatively un-weathered dark coloured mudstones of the Black Band at South Ferriby
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and Caistor; average phytoplankton is in the range of 10,000–15,000 cpg with pollen and spores at
200–900 cpg (Supplementary Table I).
Significantly higher concentrations of palynomorphs have been recorded from the dark coloured
mudstones of the Black Band and Bed G elsewhere in Lincolnshire, at Melton Ross (average
phytoplankton ca. 84,000 cpg; pollen and spores ca. 10,000 cpg; Table 2) and Bigby (average
phytoplankton ca. 60,000 cpg; pollen and spores ca. 3,500 cpg). These values may be more
representative than those from South Ferriby and Caistor, given that samples from the latter sections
are apparently more affected by quarry wall weathering. However, the relatively expanded sections at
Melton Ross and Bigby could have been deposited in local topographic troughs in the seafloor (Gaunt
et al. 1992; Wood & Mortimore 1995; Wiese et al. 2009) in which organic matter preservation may
have been enhanced (cf. Jenkyns 1985). Farther north in Yorkshire, average phytoplankton
concentrations from relatively un-weathered dark coloured mudstones of the Black Band and Bed G
are ca. 40,000–50,000 cpg at Market Weighton, Bishop Wilton and East Knapton, with pollen and
spores at ca. 3,500–5,000 cpg; i.e. values relatively comparable to Melton Ross and Bigby. The
highest concentrations of palynomorphs in the region are from sections on the margins of the
Cleveland Basin (e.g. Jeans et al. 1991; Mitchell 2000) at Flixton and Speeton, with average
phytoplankton concentrations of ca. 95,000–100,000 cpg, and pollen and spores ca. 10,000 cpg at
Speeton and ca. 80,000 cpg at Flixton (Supplementary Table I). These regional data appear to be
consistent with the trend of an increase in preservation of organic matter as the depositional
environment deepened from relatively shallow shelf to deeper continental margin (Jenkyns 1985).
Unlike the samples below the Plenus Marls at Eastbourne and Lulworth, which also contain high
concentrations of dinoflagellate cysts (ca. 38,000–75,000 cpg; see above), P. infusorioides is a
relatively minor component of assemblages from dark coloured mudstones in eastern England (Table
4), suggesting that nutrients originating from upwelling might not have been the cause of high
concentrations of dinoflagellate cysts there.
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The fact that the Wunstorf and Misburg sections are at least ten times as thick as the correlative
eastern England sections, yet show a succession of dinoflagellate cyst and acritarch bio-events
comparable to those at Melton Ross, is consistent with a high degree of stratigraphic condensation in
eastern England. The greater palynomorph concentrations documented from the dark coloured
mudstones in eastern England may partly reflect this, along with increased quality of sea floor
preservation (in a low O2 environment), particularly in potential local topographic troughs in the sea
floor and in deeper areas of the continental shelf. A loss of biogenic carbonate sediment during the
CTB mass extinction interval, and a reduction of siliciclastic input during maximum flooding
associated with the base of the Black Band (eastern England; Wood et al. 1997) and the correlative
base of the Fish Shale (NW Germany; Ernst et al. 1984), may also have contributed to stratigraphic
condensation and enhanced palynomorph concentration.
Palynomorphs are recovered from most levels within the Chalk Group of the Southern Province
(Anglo-Paris Basin; see examples above) and the Transitional Province, e.g. the Trunch borehole (Fig.
2) which has palynological recovery throughout the Upper Cretaceous section apart from a Middle
Cenomanian to Middle Turonian barren interval (Pearce 2010, 2018; Olde et al. 2015a; Pearce et al.
2020). All chalk samples analysed to date from the Cenomanian and Lower Turonian of the Northern
Province, below and above the Black Band, and from the Central Limestone at Melton Ross (this
study), are palynologically barren. This is also the case for chalk samples from the CTB interval in
NW Germany (Marshall & Batten 1988; Prauss 2006; van Helmond et al. 2015). In the North Sea
Basin, poor or patchy palynological recovery from the Chalk Group has led to palynology not being
routinely used in stratigraphic studies of hydrocarbon wells (with exceptions such as Maastrichtian
chalk in the Dan Field, Danish sector; Schiøler & Wilson 1993), whereas it is a primary stratigraphic
discipline in the coeval northerly, siliciclastic Shetland Group, which has consistent rich palynological
recovery (Costa & Davey 1992). In England, a different diagenetic history of Cenomanian and
Turonian chalks in the Northern Province, which are hardened relative to those of the Transitional and
Southern Provinces (Jeans 1980; Jeans et al. 2014), probably contributed to post-depositional loss of
palynomorphs.
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Correlation with Norwegian sector well 35/6-2 S
Ditch cuttings samples from the Grosso exploration well 35/6-2 S in the Norwegian sector, North
Viking Graben (Shetland Group facies; Figs 1, 12) were analysed for palynology by one of us (PD) in
2009. The well has a relatively thick (37 m) Blodøks Formation gamma log profile that probably
correlates with the more expanded outcrop lithological successions discussed in this paper.
A sharp rise in gamma log values marks the base of the Blodøks Formation at 2657 m (log depth).
The LO of consistent / common Litosphaeridium siphoniphorum, and rare Pterodinium
crassimuratum at 2655 m, indicate the probable presence of the pre-Plenus succession. A fall in
gamma log values from 2650.4 m to 2648.6 m may correlate with the Plenus Bed in NW Germany
and Central Limestone / Bed A in eastern England. This is supported by the FO of common
Cyclonephelium compactum–membraniphorum, Eurydinium saxoniense and Alterbidinium daveyi in
the next up-hole sample from 2646 m, in an interval of rising gamma log values from 2648.6 m to
2641.7 m (likely equivalent of eastern England Bed B). The remainder of the Blodøks Formation from
2641.7 m to 2620.0 m contains two gamma log plateaux, separated by a lower gamma interval,
probably correlative with the relatively clay- and organic-rich Fish Shale in NW Germany and Beds C
to H in eastern England. This interpretation is supported by the LO of Carpodinium obliquicostatum
from a cuttings sample near the top of the formation at 2616 m. Common C. compactum–
membraniphorum, E. saxoniense and A. daveyi persist higher than the log pick for the top of the
Blodøks Formation. The highest common occurrence of C. compactum–membraniphorum is often
used as a proxy for the top of the Lower Turonian (Dodsworth & Eldrett 2019), occurring in this well
at 2466 m. Rugubivesiculites rugosus occurs sporadically in eight samples in the Turonian interval,
from 2256 m, but is more consistent in Cenomanian deposits from 2646 m and below.
Conclusions
A sequence of dinoflagellate cyst and acritarch bio-events supports the proposed correlation of the
Melton Ross CTB succession with sections in NW Germany. The top of consistent/common
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Litosphaeridium siphoniphorum occurs in the lower succession at Melton Ross and the pre-Plenus
beds at Misburg and Wunstorf; the base of abundant Cyclonephelium compactum–membraniphorum
occurs immediately above the Central Limestone at Melton Ross, in lower Bed B, and immediately
above the Plenus Bed at Misburg and Wunstorf; a regional influx of Oligosphaeridium totum, with an
acme in upper Bed B at Melton Ross, occurs at the same level as a record of the latest Cenomanian
zonal ammonite Neocardioceras juddii at Misburg; rare specimens of the dinoflagellate cysts
Adnatosphaeridium tutulosum and Carpodinium obliquicostatum have their range tops in or just
above the Black Band in eastern England, and within the Fish Shale at Misburg and Wunstorf. An
influx of the acanthomorph acritarch Micrhystridium spp. in Bed G at Melton Ross may correlate with
that recorded from the upper parts of the Fish Shale at Wunstorf.
Correlation between eastern and southern England is less straightforward. Carbon isotope data from
Melton Ross and South Ferriby, and the distribution of dinoflagellate cyst bio-events, support the
interpretation that the Black Band is stratigraphically higher than the Plenus Marls. The restriction of
Praeactinocamax plenus (belemnite) to the Plenus Marls and Neocardioceras juddii (ammonite) to
the overlying Ballard Cliff Member are consistent with this. The correlation points to separate
depositional histories and marked differences in siliciclastic input between the Northern and Southern
provinces during post-Plenus Marls times. However, the foraminiferal and calcareous nannofossil data
can alternatively be interpreted in terms of the Black Band being equivalent to the upper part of the
Plenus Marls. Poor preservation and/or recovery of all three microfossil groups in the Ballard Cliff
Member hampers confident assessment of correlative bio-events around the CTB in southern England.
Palynological assemblages from the > 1% TOC samples at Melton Ross contain higher relative
abundances of the P-cysts Eurydinium saxoniense and Bosedinia spp., along with prasinophyte
phycomata, consistent with periodic hydrographically restricted and eutrophic conditions during
deposition of the darker mudstone layers. Exceptionally high concentrations of palynomorphs suggest
enhanced preservation in such a low O2 environment at the sediment-water interface coupled with a
high degree of stratigraphic condensation. This is in agreement with the view of Hart & Leary (1989),
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who suggested that the Black Band was deposited during ‗sluggish‘ oceanic conditions that would
have been ideal for the concentration and preservation of organic-rich sediments, with or without high
surface-water productivity.
Systematic Palaeontology
Division DINOFLAGELLATA (Bütschli, 1885) Fensome et al., 1993
Class DINOPHYCEAE Pascher, 1914
Order PERIDINIALES Haeckel, 1894
Suborder PERIDINIINEAE (Autonym)
Family PERIDINIACEAE Ehrenberg, 1831
Genus Alterbidinium Lentin & Williams, 1985
Type species: Alterbidinium “recticorne” (Vozzhennikova, 1967) Harker et al., 1990
Alterbidinium daveyi basionym nov.
Derivation of name. In honour of the palynologist Roger J. Davey.
Designation of holotype. Davey (1970, plate 1, fig. 3). Location, International Yarbo Borehole no.
17, south-east Saskatchewan, Canada (coordinates supplied by Davey, 1969, p. 115, fig. 8, are; ―east
of Regina at Lsd. 1, Sec. 24, Twp. 20, Rg. 33, W1st Meridian‖). Sample depth 254.5 m (835 ft) below
Kelly Bushing, Second White Specks Shale, Colorado Group. The slide is curated at the Natural
History Museum, London (slide/specimen reference number V.51979).
Description. See Davey (1970, p. 338).
Discussion. Davey (1970, pl. 1, figs 3, 4, p. 338) described specimens from Cenomanian deposits in
Saskatchewan that he assigned to Deflandrea (now Subtilisphaera) pirnaensis. He pointed out
differences with the type material of S. pirnaensis, as described by Alberti (1959) from Turonian
deposits in Germany, including a smaller size, length 46 (62.7) 87 μm (compared with 80–106 μm)
and width, 34 (45.5) 63 μm (compared with 58–64 μm), and presence of an archaeopyle in many
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Saskatchewan specimens but absence of one in the type specimens. It is also noted here that the
holotype of S. pirnaensis (Alberti 1959, pl. 8, fig. 1) has possible pre- and postcingular tabulation that
is absent from the Saskatchewan specimens. Stover & Evitt (1978) proposed a new species, Alterbia
daveyi, based on the specimens illustrated in Davey (1970) but did not designate one of the specimens
as a holotype. The name was therefore not validly published (Fensome et al. 2019). Subsequently, the
genus Alterbia has been considered an illegitimate name and its species have been transferred to the
genus Alterbidinium (Fensome et al. 2019). Alterbidinium “daveyi” has become widely accepted as a
separate but informal species. The designation herein of one of the specimens illustrated in Davey
(1970) as a holotype, gives it formal status.
Alterbidinium daveyi and Eurydinium saxoniense have comparable morphology, and both possess a
single intercalary (2a) plate archaeopyle which is steno- to iso-deltaform. Alterbidinium daveyi may
be slightly larger (cf. length, 52–66 μm, and width, 37–54 μm for E. saxoniense; Marshall & Batten
1988) and appears to have a more strongly-developed apical horn and cingulum than E. saxoniense,
but intergradations may occur. The cingulum in A. daveyi is marked by low ridges that sometimes
possess pustules distally and is occasionally crossed by low ridges delimiting plate boundaries (Davey
1970, p. 338); the latter features are not reported in E. saxoniense. The holotype of A. daveyi appears
to show some anterior dorsal intercalary tabulation (2a and 3a plate boundaries). Dorsal tabulation is
usually restricted to the archaeopyle in E. saxoniense (Marshall & Batten 1988), though our illustrated
specimen (Fig. 5.6) may possess sutures around both 2a and 3a intercalary plates.
The eastern England specimens inspected in this paper and previous studies have been assigned to E.
saxoniense, although the additional presence of A. daveyi cannot be ruled out. The dinoflagellate cyst
distribution charts of Marshall & Batten (1988) indicate an absence of A. daveyi and the common to
abundant occurrence of E. saxoniense from the (post-Plenus) CTB interval in NW Germany.
Conversely, in coeval deposits from North America, A. daveyi (sometimes recorded as S. pirnaensis)
is prominent (Bloch et al. 1999, fig. 23; Dodsworth 2000, 2016; Harris & Tocher 2003; Dodsworth &
Eldrett 2019). In the Shetland Group of Northern Europe, both A. daveyi and E. saxoniense are
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common, e.g. in Norwegian well 35/6-2 S, where confident differentiation between the species can be
problematic for some specimens.
Genus Bosedinia He, 1984, emend. Chen et al., 1988, Prauss, 2012c
Type species: Bosedinia granulata (He & Qian, 1979) He, 1984
Bosedinia laevigata (Jiabo, 1978, ex He & Qian, 1979) He, 1984
Figs 7.11, 7.12, 7.13, 7.17, 7.21
Description. Small to intermediate-sized, sub-spheroidal shaped, smooth and thin-walled proximate
autocysts with an ornament of ca. 10–50 sparsely distributed, non-tabular, solid verrucae, each of
which is ca. 1–3 μm in diameter. Tabulation is indicated by an archaeopyle only. A flap-like
operculum, probably involving fused apical and anterior intercalary plates, is often attached.
Inclusions (omphali) are present in some specimens.
Dimensions. Diameter (central body, 12 measured specimens): 35.7 (43.5) 56.0 µm.
Discussion. The type specimens of Bosedinia laevigata (He et al. 2009, pl. 17, figs 11–15), from
lacustrine Neogene deposits in China, are of comparable size (35–60 μm diameter) to the specimens
measured from Melton Ross, and contain a comparable number, size and distribution of small
verrucae ornament. Bosedinia laevigata has been recorded from marine Upper Cretaceous deposits at
Tarfaya, Morocco (Prauss 2012c, 2015) and the Abu Gharadig Basin in Egypt (Ahmed et al. 2020;
Fig. 1), but not previously in Europe.
Stratigraphic range/occurrence. At Melton Ross, Bed C (sample -11.5) to Bed E (sample -8).
Bosedinia cf. sp. 1 of Prauss (2012b)
Figs 7.5, 7.9, 7.10
Discussion. The abundant occurrence of mainly enclosed spheres, containing omphali, in organic-rich
Cenomanian–Coniacian deposits at Tarfaya, has been discussed by Prauss (2012a, b, c, 2015). Prauss
(2012b) provided evidence for their assignment to the genus Bosedinia from occasional specimens
that have an archaeopyle/operculum. Eldrett et al. (2017) documented the abundant occurrence of
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comparable specimens at other proto-Atlantic sites (Demerara Rise, DSDP Sites 1260 and 1261; Fig.
1) and within the Western Interior Seaway of the United States (SW Texas and central Colorado).
Specimens from Melton Ross constitute the first record of their presence in Europe. However, there
are issues with consistent identification. While the specimen illustrated in Fig. 7.9 shows a rare
example of discernible anterior intercalary and apical plates being involved in operculum formation,
consistent with Bosedinia, the fully enclosed specimen in Fig. 7.10 could alternatively be interpreted
as an endocyst of Subtilisphaera pontis-mariae. (cf. the specimens in Figs 5.14, 5.15). The two
overlapping specimens in Fig. 7.5, resemble Eyrea nebulosa, which also possesses omphali (cf. plate
11, in Cookson & Eisenack 1971), but may be distinguished by the presence of a kalyptra, when
preserved. In the Melton Ross data, specimens encountered are mainly single enclosed spheres with
omphali and have been assigned to ―Bosedinia cf. sp. 1 / peridinioid endocysts‖ in the supplementary
distribution chart.
Stratigraphic range/occurrence. At Melton Ross, local Bed II (sample -21) to Bed H (sample -1).
Genus Ginginodinium Cookson & Eisenack, 1960, emend. Lentin & Williams, 1976
Type species: Ginginodinium spinulosum Cookson and Eisenack, 1960
Ginginodinium? sp. A of Prauss, 2006, 2012a
Figs 6.3, 6.7
Discussion. In Cenomanian and Turonian deposits from NW Germany (Wunstorf) and Morocco
(Tarfaya), Prauss (2006, 2012a) differentiated specimens that are morphologically close to
Trithyrodinium suspectum but with a more complex archaeopyle, tentatively assigning them to the
genus Ginginodinium. The archaeopyle in Ginginodinium? sp. A involves from one intercalary plate
(2a) to three intercalary plates (1a–3a), often attached, and sutures are usually present between
precingular plates which are attached at the cingular border (3I3Pa). Prauss (2006) noted that
Ginginodinium? sp. A probably includes specimens informally described as ‗Cyst form A‘ by
Marshall & Batten (1988). The type species, Ginginodinium spinulosum (Cookson & Eisenack 1960,
p. 7, pl. 2, fig. 9), and other accepted species, sometimes exhibit accessory sutures on dorsal
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precingular plates, in addition to a three plate intercalary archaeopyle (Stover & Evitt 1978). In the
Melton Ross material, consistent differentiation of this taxon from T. suspectum was problematic.
Stratigraphic range/occurrence. At Melton Ross, local Bed I (sample -22) to Bed H (sample -1).
Genus Subtilisphaera Jain & Millepied, 1973, emend. Lentin & Williams, 1976
Type species: Subtilisphaera senegalensis Jain & Millepied, 1973
Subtilisphaera pontis-mariae (Deflandre, 1936) Lentin & Williams, 1976
Figs 5.14, 5.15
Description. Small to intermediate-sized, elongate to ovoidal-shaped, smooth-walled, bicavate to
circumcavate peridinioid dinoflagellate cysts with pericoels developed into a sub-conical apical horn
and one similar antapical horn which is positioned asymmetrically. The epicyst is larger than the
hypocyst. The cingulum is delimited by two low ridges. An archaeopyle has not been observed. The
endocyst is sub-spherical and slightly thicker walled than the periphragm. The horns are
approximately 1/5 to 1/4 of the diameter of the endocyst.
Dimensions (seven measured specimens). Length 47.3 (52.7) 57.0 µm; width 32.6 (36.0) 42.2 µm.
Discussion. The Subtilisphaera specimens encountered in the present study have comparable
morphology, with some variation in the degree of cavation and the length of horns. They are recorded
here as Subtilisphaera pontis-mariae. The holotype (Deflandre 1936, pl. 2, fig.7) is closely
comparable to the Melton Ross specimen illustrated in Fig. 5.14. Some subsequently illustrated
specimens (e.g. Davey 1970, pl.1, figs 10–11; Dodsworth 2016, pl. 1, fig. 7) differ slightly in having
longer horns, ca. 1/2 to 1/3 the diameter of the endocyst. While Dodsworth (1996, 2000) used
Subtilisphaera spp. for eastern and southern England specimens from the CTB interval, other workers
in eastern England (Marshall & Batten 1988; Hart et al. 1993), southern England (FitzPatrick 1995;
Pearce et al. 2009) and northern France (Foucher 1979) have considered the range of morphological
variation of Subtilisphaera specimens to fall within acceptable intra-specific limits for S. pontis-
mariae in material of this age. However, Foucher (1980) assigned latest Cenomanian specimens to S.
cf. pontis-mariae in the Boulonnais region of northern France.
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Stratigraphic range/occurrence. At Melton Ross, local Bed I (sample -22) to Bed H (sample -1).
Genus Trithyrodinium Drugg, 1967, emend. Davey, 1969, Lentin & Williams, 1976, Marheinecke,
1992
Type species: Trithyrodinium evittii Drugg, 1967
Trithyrodinium maculatum sp. nov.
Figs 7.14, 7.15, 7.18, 7.19
Derivation of name. From maculate, meaning spotted or stained, with reference to the distinctive
surface markings.
Designation of holotype. Fig. 7.18. Sample MR3. Slide MR3(A). Coordinates S32/3. Location,
Melton Ross Quarry, Lincolnshire, UK. Stratum, Bed F, Flixton Member, Welton Chalk Formation.
The specimen is curated in the MPK collection of the British Geological Survey, Keyworth,
Nottingham, U.K., specimen number MPK 14662.
Diagnosis. A species of Trithyrodinium possessing an ornament of ring-shaped indentations on the
endophragm and periphragm.
Description. Small to intermediate-sized, spheroidal to ovoidal-shaped, circumcavate, peridinioid
dinoflagellate cysts. The endophragm (ca. 1 μm thick) and periphragm (ca. 0.5 μm thick) are smooth
to finely-granular, both possessing regular ornament elements that are circular to sub-circular, ring-
shaped indentations (ca. 3–6 μm diameter), which are at least half the thickness of the surrounding
wall areas. Cavation is ca. 2–3 μm wide at the lateral margins and up to 10 μm in the antapical region.
An archaeopyle involving three anterior intercalary plates, type 3I(1–3a), is formed in the endocyst, with
operculum plates attached or detached. The 2a intercalary plate is isodeltaform, hexa-type (Fig. 7.19;
cf. text-figure 6 in Bujak & Davies 1983). The periarchaeopyle type has not been confirmed. There is
no evidence of tabulation, other than that indicated by the archaeopyle.
Dimensions (length x width of the central endocyst). Holotype, 39.6 x 42.5 µm. Other specimens (5
measured): length 38.0 (50.0) 66.9 µm, width 39.5 (45.6) 55.3 µm.
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Discussion. Only six specimens of this taxon have been recorded to date. However, its distinctive
morphology makes it easy to identify and warrants the erection of a formal species. The endocyst
archaeopyle type is consistent with the genus Trithyrodinium but an outer wall layer (periphragm) has
only been observed on two specimens, including the holotype. The circular indentations on the cyst
walls are ring-shaped, as opposed to the whole area within the circle being thinned. The ring shapes
are reminiscent of those attributed to impressions made by coccoliths on the surface of specimens of
an alga (Campenia sp.) by Prauss (2012b, figs 10A, 11H). However, the T. maculatum specimens
occur in assemblages of abundant dinoflagellate cysts, with other taxa not exhibiting comparable
markings. The markings are a key identifying feature but could be a preservation artefact. This type of
surface ornament has not previously been reported on other species of P-cysts, including those
belonging to the genus Trithyrodinium. In the G-cyst species Apteodinium maculatum, comparable
ring-shaped thinning surrounds small thickened circular areas (Eisenack & Cookson 1960).
Stratigraphic range/occurrence. At Melton Ross, Bed B (sample -12) to Bed F (sample -3).
Trithyrodinium? sp. A (this paper)
Figs 7.1, 7.2
Description. Small to intermediate-sized, sub-polygonal to peridinioid-shaped, smooth and thin-
walled cysts that lack ornament. Peridinioid tabulation is indicated by sutures or breaks between some
plates on the epicyst, indicating three anterior intercalary plates. An archaeopyle has not been
observed. A periphragm has not been observed. There is no evidence of tabulation, other than
indicated by the sutures.
Dimensions (length x width). First specimen, 46.5 x 40.7 µm; second specimen, 54.3 x 50.0 µm.
Discussion. Prauss (2012a, fig. 15) illustrated an unpublished smooth, thin-walled species of
Trithyrodinium at Tarfaya that usually has sutures around three intercalary plates and rarely possesses
a periphragm (Prauss, 2012a, fig. 15J). It is uncertain whether this taxon is the same as the Melton
Ross specimens described here.
Stratigraphic range/occurrence. At Melton Ross, Bed C (sample -11) to Bed G (sample -2).
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Order GONYAULACALES Taylor, 1980
Suborder GONYAULACINEAE (Autonym)
Family GONYAULACACEAE Lindemann, 1928
Genus Dissiliodinium Drugg, 1978
Type species: Dissiliodinium globulus Drugg, 1978
?Dissiliodinium globulus Drugg, 1978
Figs 7.4, 7.8
Description. Intermediate-sized, spheroidal-shaped, smooth and thin-walled proximate autocysts
lacking ornament. Tabulation is indicated by sutures between precingular plates on the epicyst.
Archaeopyle formation involves one or more precingular plate. There is no evidence of tabulation on
the hypocyst.
Dimensions (four specimens measured). Diameter: 57.0 (60.7) 65 µm.
Discussion. The observed features of the four available specimens allow tentative assignment to
Dissiliodinium globulus. The taxon has been documented from the Upper Jurassic of Central Europe
(Drugg 1978) and Northern Europe (e.g. Bailey et al. 1997) and is present in higher proportions
within the Lower Cretaceous of SW Morocco (Below 1981). Prauss (2012b, fig 9a–i) reported
specimens from around the CTB at Tarfaya.
Stratigraphic range/occurrence. At Melton Ross, Bed E (sample -8 only).
Genus Leptodinium Klement, 1960, emend. Stover & Evitt, 1978, Sarjeant, 1982
Type species: Leptodinium subtile Klement, 1960
Leptodinium? aff. delicatum (this paper)
Fig. 6.12
Description. Intermediate-sized, smooth and thin-walled autocysts with sub-polygonal shape and
septae (entire; ca. 1.5 to 3.5 μm high) which probably correspond to tabulation. The cingulum appears
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to have strongly offset ventral ends. An archaeopyle has not been observed on the two specimens
recorded.
Dimensions. Diameter: illustrated specimen, 61.8 µm; second specimen, 60.8 µm.
Discussion. In Leptodinium? delicatum, the wall is very thin (less than 0.5 μm thick) and only attains
a thickness of 0.5 μm when forming plate boundaries (Davey 1969, p. 123).The Melton Ross
specimens resemble L.? delicatum but differ in possessing sutural septae. The detailed plate formula is
not yet determined. The probable offset sulcus is consistent with the genus Leptodinium though the
presence of septae is more compatible with the genus Impagidinium (Stover & Evitt 1978).
Stratigraphic range/occurrence. At Melton Ross, Bed C (samples -11 and -11.5).
Genus Oligosphaeridium Davey & Williams, 1966, emend. Davey, 1982
Type species: Oligosphaeridium complex (White, 1842) Davey & Williams, 1966
Oligosphaeridium totum Brideaux, 1971
Figs 5.9, 5.10, 8.9
Discussion. The published taxon Oligosphaeridium totum is considered here to accommodate a
skolochorate dinoflagellate cyst from NW Germany and the North Sea that was informally assigned
by Marshall & Batten (1988) to Litosphaeridium sp. A. The latter name was adopted in subsequent
studies of the CTB interval in eastern England (Hart et al. 1993; Dodsworth 1996), southern England
(FitzPatrick 1995) and Poland (Dodsworth 2004b). The processes of O. totum are tubular, sometimes
slightly narrower medially, with apices usually flaring, buccinate and possessing an entire margin
(Brideaux 1971). Two subspecies are differentiated, based mainly on process length relative to central
body, O. totum subsp. totum (relatively long processes, approximately two-thirds to one central body
diameter in length; comparable to most specimens encountered in this study, e.g. Figs 5.9, 8.9) and O.
totum subsp. minus (relatively short processes, approximately one half of the central body diameter;
cf. Fig. 5.10). The Melton Ross specimens have the same process formula as those described by
Brideaux (1971) for O. totum and by Marshall & Batten (1988) for Litosphaeridium sp. A; 4I, 6II, 0c,
5III, 1p, 1IIII, 1s. This is a process formula common to all species of Oligosphaeridium and some
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species of Litosphaeridium (Lucas-Clark 1984). Litosphaeridium differs from Oligosphaeridium in
having dome-shaped processes that are typically not expanded distally (Stover & Evitt 1978). The
type material of O. totum has a slightly thicker, more scabrate central body wall than the smooth walls
observed on Melton Ross specimens and those described and illustrated from NW Germany (Marshall
& Batten 1988, p. 92, pl. I, fig. 9; Marshall 1983, pl. 13, figs 16, 18). However, Singh (1971) reported
a smooth cyst wall on Oligosphaeridium diastema, a junior synonym of O. totum (as agreed by Singh
and Brideaux; Brideaux & McIntyre 1975, p. 29). Some variation in thickness and ornament of the
central body may therefore be attributed to intra-specific variation.
Stratigraphic range/occurrence. At Melton Ross, local Bed VII (sample -17) to Bed F (sample -3).
Genus Pterodinium Eisenack, 1958, emend. Yun, 1981, Sarjeant, 1985
Type species: Pterodinium aliferum Eisenack, 1958
Pterodinium crassimuratum (Davey & Williams, 1966) Thurow et al., 1988
Fig. 5.7
Discussion. Kjellström (1973) and Pavlishina (1990) considered Pterodinium? pterotum to be a senior
synonym of Pterodinium crassimuratum and Pterodinium cingulatum subsp. polygonale. In
Dodsworth & Eldrett (2019, plate 1, fig.4), specimens encountered were assigned on the basis of this
proposed synonymy to P.? pterotum. However, subsequent inspection of the holotype photographs
and original descriptions for the three taxa indicates that our specimens, both from the United States
and Europe, are closest to P. crassimuratum and P. cingulatum polygonale, having thickened intra-
plate areas and ―a linear depressed area on each side of the ledges [septae] thereby separating a more
elevated central plate area from the ledges‖ (Clarke & Verdier 1967, p. 47). We agree with Clarke et
al. (1968, p. 181) that the latter subspecies is a junior synonym of P. crassimuratum. However,
Cookson & Eisenack (1958, p. 50) make no reference to comparable thickened intra-plate areas in P.?
pterotum, and none are visible on their holotype photograph (plate 11, fig. 7). We therefore conclude
that P.? pterotum is not a senior synonym of P. crassimuratum.
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Family AREOLIGERACEAE Evitt, 1963
Genus Aptea Eisenack, 1958
Type species: Aptea polymorpha Eisenack, 1958
Aptea? spongireticulata (Prössl, 1990 ex Prössl, 1992) Fensome et al., 2019
Figs 6.1, 6.2
Description. Intermediate-sized, lenticular-shaped cysts with rounded or flattened apical and
antapical areas, lacking well-developed horns. The autophragm is smooth to lightly pitted and is
covered with a coarse reticulum. Rod-like structures with expanded to bifurcating tips support the
reticulum muri and are ca. 6–10 μm long. The muri are fibrous or finely fenestrate. The lacuna
between the muri vary in size, from ca. 5–13 μm across. A slight indentation in the reticulum in the
antapical region on one specimen (Fig. 6.1) may represent a gap between antapical horns. Tabulation
is indicated by an apical archaeopyle only.
Dimensions (five measured specimens, length x width without operculum). Length 61.0 (66.9) 73.0
µm; width 83.8 (86.3) 89.0 µm.
Discussion. The Melton Ross specimens conform to the type material of A.? spongireticulata,
illustrated from NW Germany by Prössl (1990). Canningia macroreticulata differs in having smaller
lacuna between its muri of ca. 2–7 μm and has shorter (ca. 2 μm high), entire muri without
fibrous/fenestrate structure (Lebedeva, in Ilyina et al. 1994). The Melton Ross specimens may be the
first published record of A.? spongireticulata outside Germany.
Stratigraphic range/occurrence. At Melton Ross, Bed B (sample -13) to Bed F (sample -5). In NW
Germany, Prössl (1990) reported an Albian to Middle Turonian range for A.? spongireticulata.
Genus Canninginopsis Cookson & Eisenack, 1962
Type species: Canninginopsis denticulata Cookson & Eisenack, 1962
Canninginopsis? lindseyensis sp. nov.
Figs 6.8, 6.9, 7.7
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Derivation of name. From Lindsey, a former Anglo-Saxon kingdom and current district of
Lincolnshire.
Designation of holotype. Figures 6.8, 6.9. Sample MR11. Slide MR11(B). Coordinates P44/3.
Location, Melton Ross Quarry, Lincolnshire, UK. Stratum, Bed C, Flixton Member, Welton Chalk
Formation. The specimen is curated in the MPK collection of the British Geological Survey,
Keyworth, Nottingham, U.K., specimen number MPK 14663.
Diagnosis. An areoligeracean dinoflagellate cyst possessing tabular to penitabular thickenings that
form the bases of thinner, entire septae.
Description. Small to intermediate-sized, lenticular-shaped autocysts with differential development of
a larger left antapical horn. A sulcal notch is present. The cyst wall is ca. 1 μm thick and has a smooth
to slightly reticulate surface. Tabulation is indicated by an apical archaeopyle (operculum detached)
and tabular to penitabular thickenings, ca. 1–2 μm high and 1–3 μm wide, that form the bases of
thinner septae. The septae are ca. 2–5 μm high and 1 μm wide, entire, smooth to slightly reticulate and
often exhibit some longitudinal folding (‗creases‘). Proximal thickenings/septae are adjacent (tabular)
or separated by gaps of up to 4 μm (penitabular). The gonyaulacoid tabulation pattern is (operculum
not observed); 6II, 6c, 5–6III, ?1P, 1IIII, ?s.
Dimensions (length x width, without operculum). Holotype: length 44.4 µm, width 54.1 µm. Other
specimens (2 measured): length 43.0 (46.3) 49.6 µm, width 58.9 (64.3) 69.6 µm.
Discussion. The shape, presence of a sulcal notch and a probable small postcingular 1III plate (Fig.
6.8) indicate an areoligeracean affinity (cf. Evitt 1985, text-figs 10.2, 10.6). Strongly developed
tabulation is consistent with the genus Canninginopsis. Although the type species and most other
accepted species often possess discontinuous ornament elements reflecting tabulation such as spines
or grana, continuous septae are present on the species Canninginopsis maastrichtiensis from the
Maastrichtian of Belgium and Netherlands (Slimani 1994). However, the development of gaps
between septae, giving rise to penitabular ornament, is inconsistent with Canninginopsis and warrants
questionable assignment to the genus. Penitabulation has been noted in the related genus Canningia,
e.g. Canningia transitoria (Stover & Helby 1987, fig. 4) but its septae are coarsely perforate.
Schematophora possesses entire to occasionally perforate penitabular ridges but has a spherical, non-
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areoligeracean shape and lacks any tabulation in the cingulum region (Deflandre & Cookson 1955;
Stover & Evitt 1978). On the holotype of C.? lindseyensis, the dorsal precingular plate (3II) appears to
be absent but this could be due to specimen damage. Likewise, a precingular 6II plate may be
damaged. A second specimen with less well-developed or preserved septae (Fig. 7.7) indicates that six
precingular plates may usually be present. The highly distinctive morphology warrants erection of a
formal species, although it is acknowledged that doing so is unusual for just three specimens.
Recovery and inspection of further specimens is required to fully elucidate the taxon and its generic
assignment.
Stratigraphic range/occurrence. At Melton Ross, Bed C (sample -11 only).
Dinoflagellate cysts of uncertain supra-generic affinity
Dinoflagellate? type D of Ioannides, 1986
Figs 6.13, 6.14
Dimensions (27 specimens). Diameter (autophragm central body): 45.7 (60.7) 75.2 µm.
Discussion. Ioannides (1986, p. 42, pl. 25, figs 1– 4, 6) described comparable though slightly larger
taxa from Upper Cretaceous (Santonian–Maastrichtian) deposits in Arctic Canada (size range 70–79
μm length and 70–87 μm width; 11 measured specimens). He noted that ornament varied from
granular to minutely verrucate/rugulate, and that, ―in some specimens, a number of ‗slits‘ have been
observed along the autophragm. Although a weak plate attachment may be suggested, no definite
regular pattern has been determined. Occasionally, six paraplates (precingular or postcingular) may be
visualised‖, as shown here in Figs 6.13, 6.14. A similar but larger form (140 x 119 μm), surrounded
by a kalyptra, was described from Western Australia by Cookson & Eisenack (1971, p. 223, plate 11,
fig. 1) as Eyrea sp. Occasional Melton Ross specimens possess a kalyptra, comparable to Eyrea sp.,
but most specimens lack an outer layer. Comparable specimens from Poland were assigned to Eyrea?
spp. by Dodsworth (2004b). In poorly preserved material from eastern England, it can be difficult to
distinguish specimens of Dinoflagellate? type D from degraded and/or broken specimens of
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Sentusidinium ringnesiorum (cf. Fig. 7.20) and the granular endocysts of Trithyrodinium suspectum
and Ginginodinium? sp. A.
Stratigraphic range/occurrence. At Melton Ross, local Bed I to Bed H. A broad Late Cretaceous
range is suggested by the records of Ioannides (1986) and Cookson & Eisenack (1971).
Acknowledgements
Richard Stansfield, CEO of the Singleton Birch Group, is thanked for allowing access to Melton Ross
Quarry in 1997 and encouraging publication of new data. In 2019, quarry staff members Ben Hyde
and Dave Alliss provided an update on the sampled sites. PD undertook palynological laboratory
processing of the Melton Ross samples at the former Aberdeen office of Robertson Research
International Ltd while employed there as a biostratigrapher in 1997. The company and its former
management, Keith L. Marshall and Grant Heath, are thanked for providing the laboratory facilities.
KLM is also thanked for permission to refer to his unpublished PhD thesis. Tim Absalom and James
Quinn of Plymouth University drafted Figure 2. Jennie Houlgrave of RPS Ichron prepared Figure 3
and assisted with Figure 4. Natalia Lebedeva, Klaus Prössl and staff at the Geological Society library
assisted with gathering literature. Organic geochemistry and stable isotope work were commissioned
by StrataSolve Ltd in 2019. TOC and Rock-Eval pyrolysis analyses were undertaken by Adam
Fermor and Patrick Barnard of APT (U.K.) Ltd. Stable isotope analyses were conducted by
Christopher Day, Department of Earth Sciences, Oxford University. StatoilHydro AS (now Equinor
Energy AS) are thanked for permission in 2010 to publish the CTB section from their Norwegian well
35/6-2 S. James P. Fenton and Martin A. Pearce (reviewers), and Stewart G. Molyneux (editor) made
helpful suggestions for improving the manuscript.
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Table captions
Table 1. Lowermost part of the Welton Chalk Formation: A comparison of bed nomenclature. Sample
positions are indicated.
Table 2. Summary of palynomorph concentration (counts per gramme), total organic carbon (TOC),
palynomorph groups (gymnosperm pollen; pteridophyte/bryophyte spores; angiosperm pollen;
dinoflagellate cysts; prasinophyte phycomata; acanthomorph acritarchs), separate > 10 µm kerogen
counts of 100 particles (AOM = amorphous organic matter; phytoclasts; palynomorphs) and
dinoflagellate cyst species richness (‗diversity‘). Shaded rows highlight samples with > 1 wt.% TOC.
T/M ratio = terrestrial/marine ratio. P/G ratio = the ratio between peridinioid (P) and gonyaulacoid
(G) dinoflagellate cysts.
Table 3. Relative and absolute abundance of selected gonyaulacoid dinoflagellate cysts (G-cysts), and
Dinoflagellate? type D of Ioannides (1986). P = Present, not quantified (rare occurrence, outside the
300 palynomorph count). The calculation of absolute abundance per gram of sample (cpg) is
explained in the text.
Table 4. Relative and absolute abundance of common/abundant peridinioid dinoflagellate cysts (P-
cysts). P = Present, not quantified (rare occurrence, outside the 300 palynomorph count). The
calculation of absolute abundance per gram of sample (cpg) is explained in the text.
Figure captions
Fig. 1. Turonian palaeogeographic reconstruction with the main Northern Hemisphere site locations
discussed in the text: brown shaded area, landmass; light blue, palaeo-shelf; CLIP, Caribbean large
igneous province; HALIP, high-Arctic large igneous province. Modified from Eldrett et al. (2014,
2017) and Du Vivier et al. (2015).
Fig. 2. The Cretaceous outcrop in eastern England and the location of sections discussed in the text
(modified from Hart 2019). The Black Band wedges out to the south of Louth (line A–A). It appears
to represent the ‗feather-edge‘ of OAE-2 when traced towards a palaeo-high located in the region of
the Wash. Between lines A–A and B–B the chalks and nodular chalks of latest Cenomanian – earliest
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Turonian age are dull red in colour, while south of line B–B the chalks at the same stratigraphic level
are pale green/grey in colour.
Fig. 3. Sketch map of Melton Ross Quarry, north Lincolnshire, showing the position of the
Cenomanian–Turonian boundary (CTB) succession excavations in 1997 (after Wood et al. 1997). Site
1 and Site 4 sampling exposures (located around 53º 35‘ 14‖ N, 0º 22‘ 32‖ W at 19 m above sea level)
are currently (2019) buried approximately 10 m below the restored surface of the quarry.
Fig. 4. Composite lithological log of the sampled Melton Ross Quarry section with carbon and
oxygen isotope curves. Biostratigraphy: 1, planktonic foraminiferal zones extrapolated from Elsham
and South Ferriby (Hart & Bigg 1981; Hart & Leary 1989); 2, 3, dinoflagellate cyst zones and
subzone of Olde et al. (2015a) and Dodsworth & Eldrett (2019). Formations of Wood & Smith
(1978). Members of Jeans (1980), Mitchell (2000) and Hopson (2005; new sub-member rank). The
Buckton Member is ‗The Inoceramite‘ of Hart et al. (1991, 1993) and Hart (2019). Local beds I–VII
of Wood et al. (1997), regional Beds A–H of Dodsworth (1996), Turnus Bed of Pomerol &
Mortimore (1993), ‗Adrian‘s Pair of Marls‘ of Mortimore (2014), Central Limestone of Wood et al.
(1997) and Hildreth (1999), Black Band sensu Wood & Smith (1978). Distribution of the belemnite
Praeactinocamax plenus was reported from Site 1 at Melton Ross by Wood et al. (1997). OAE-2 is
picked based on relatively high δ13C values of greater than 3‰ VPDB in this section. Points ‗a‘, ‗b‘
and ‗c‘ are tentatively correlated with the corresponding southern England picks of Jarvis et al.
(2006), allowing tentative positioning of CTB around the Bed F/G boundary.
Fig. 5. Dinoflagellate cysts and miospores from Melton Ross. All specimens were photographed at x
400. Sample/slide numbers, specimen England Finder co-ordinates, and BGS specimen registration
numbers (MPK prefix) are given. Specimens 1–16 are dinoflagellate cysts; 17–19 are gymnosperm
pollen; 20–21 are angiosperm (Normapolles) pollen. The 50 μm scale bar applies to all specimens.
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1. Litosphaeridium siphoniphorum (Cookson & Eisenack, 1958) Davey & Williams, 1966, local Bed
I, MR97-22(A), H43/0, MPK 14664;
2. Wrevittia cassidata (Eisenack & Cookson, 1960) Helenes & Lucas-Clark, 1997, local Bed II,
MR97-21(A), S43/3, MPK 14665;
3. Carpodinium obliquicostatum Cookson & Hughes, 1964, Bed C, MR97-11.5(B), X47/1, MPK
14666;
4. Adnatosphaeridium tutulosum (Cookson & Eisenack, 1960) Morgan, 1980, local Bed II, MR97-
21(B), W31/2, MPK 14667;
5. Cyclonephelium compactum Deflandre & Cookson, 1955 – Cyclonephelium membraniphorum
Cookson & Eisenack, 1962, transitional ‗complex‘ of Marshall & Batten (1988), Bed C, MR97-
11.5(D), Q36/2, MPK 14668;
6. Eurydinium saxoniense Marshall & Batten, 1988, Bed E, MR97-7(B), J42/4, MPK 14669;
7. Pterodinium crassimuratum (Davey & Williams, 1966) Thurow et al., 1988, local Bed I, MR97-
22(B), F47/3, MPK 14670;
8. Adnatosphaeridium? chonetum (Cookson & Eisenack, 1962) Davey, 1969, local Bed II, MR97-
21(A), S38/3, MPK 14671;
9. Oligosphaeridium totum Brideaux, 1971, Bed C, MR97-11.5(A), F36/0, MPK 14672. This taxon
was recorded as Litosphaeridium sp. A by Marshall & Batten (1988);
10. O. totum, Bed C, MR97-11.5(A), W42/0, MPK 14673;
11. Stephodinium coronatum Deflandre, 1936, Bed E, MR97-8.5(D), V42/2, MPK 14674;
12. Trithyrodinium suspectum (Manum & Cookson, 1964) Davey, 1969, Bed E, MR97-8(A), T35/2,
MPK 14675;
13. Palaeohystrichophora infusorioides Deflandre, 1935, local Bed II, MR97-21(A), Q28/4, MPK
14676;
14. Subtilisphaera pontis-mariae (Deflandre, 1936) Lentin & Williams, 1976, Bed C, MR97-11.5(D),
R42/1, MPK 14677;
15. S. pontis-mariae, Bed E, MR97-8.5(A), Q25/2, MPK 14678;
16. Sepispinula? huguoniotii (Valensi, 1955) Islam, 1993, Bed C, MR97-10(A), P47/0, MPK 14679;
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17. Rugubivesiculites rugosus Pierce, 1961, Bed E, MR97-8(A), R21/1, MPK 14680;
18. Alisporites microsaccus (Couper, 1958) Pocock, 1962, Bed C, MR97-11.5(A), F38/1, MPK
14681;
19. Classopollis spp., Bed G, MR97-2(C), P36/2, MPK 14682;
20. Atlantopollis microreticulatus Krutzsch, 1967, Bed D, MR97-10.5(C), E35/3, MPK 14683;
21. Complexiopollis spp., Bed E, MR97-6(C), N41/4, MPK 14684.
Fig. 6. Dinoflagellate cysts from Melton Ross. All specimens were photographed at x 400.
Sample/slide numbers, specimen England Finder co-ordinates, and BGS specimen registration
numbers (MPK prefix) are given. The 50 μm scale bar applies to all specimens.
1. Aptea? spongireticulata (Prössl, 1990, ex Prössl, 1992) Fensome et al., 2019, Bed B, MR97-12(D),
E32/3, MPK 14685;
2. A.? spongireticulata, Bed D, MR97-9(A), P37/1, MPK 14686;
3. Ginginodinium? sp. A of Prauss (2006, 2012a), Bed D, MR97-10(C), T42/3, MPK 14687. The
arrow indicates a probable suture between two precingular plates;
4. Cyclonephelium membraniphorum Cookson & Eisenack, 1962, Bed E, MR97-8.5(C), U28/0, MPK
14688;
5. Heslertonia striata (Eisenack & Cookson, 1960) Norvick, 1976, Bed H, MR97-1(D), U45/0, MPK
14689;
6. Microdinium setosum Sarjeant, 1966, Bed C, MR97-11(D), K47/3, MPK 14690;
7. Ginginodinium? sp. A, Bed E, MR97-8.5(A), Q29/4, MPK 14691. The arrow indicates a probable
suture between two precingular plates;
8–9. Canninginopsis? lindseyensis sp. nov., Bed C, MR97-11(B), P44/3, MPK 14663. Holotype. 8.
Focus on ventral surface. Note the interpretation of tabulation and sulcal notch (arrow). 9. Same
specimen, focus on distal surface;
10. Ellipsodinium rugulosum Clarke & Verdier, 1967, local Bed II, MR97-21(A), N34/2, MPK
14692;
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11. Ginginodinium? sp. A, or an endocyst of Trithyrodinium suspectum, Bed H, MR97-1(C), V29/0,
MPK 14693;
12. Leptodinium? aff. delicata (this paper), Bed C, MR97-11.5(C), R38/1, MPK 14694. The arrows
indicate the possible lateral position of a cingulum;
13. Dinoflagellate? type D of Ioannides (1986), Bed C, MR97-8(B), U29/0, MPK 14695. Note the
presence of three or four possible precingular plates and two intercalary or apical plates on the upper
part of the cyst;
14. Dinoflagellate? type D, local Bed II, MR97-20(A), S47/4, MPK 14696. Note the presence of at
least four ‗plates‘ that resemble a precingular series on the upper part of the cyst and further sutures
on the lower part of the cyst.
Fig. 7. Dinoflagellate cysts from Melton Ross. All specimens were photographed at x 400.
Sample/slide numbers, specimen England Finder co-ordinates, and BGS specimen registration
numbers (MPK prefix) are given. The 50 μm scale bar applies to all specimens.
1. Trithyrodinium? sp. A (this paper), Bed E, MR97-8(C), W29/0, MPK 14697. Note the sub-
polygonal/peridinioid shape, laevigate surface and tabulation interpretation, including a prominent
suture/break above the three intercalary plates;
2. Trithyrodinium? sp. A, Bed E, MR97-8(A), R32/2, MPK 14698. Note the sub-
polygonal/peridinioid shape, tabular sutures and some probable breakage in the upper-right part of the
cyst;
3. Kalyptea spp., Bed C, MR97-11.5(C), V30/0, MPK 14699;
4. ?Dissiliodinium globulus Drugg, 1978, Bed E, MR97-8(C), N39/4, MPK 14700. Polar–dorsal
epicystal compression with sutures between some dorsal precingular plates, plus a possible one plate
archaeopyle with attached operculum;
5. Bosedinia cf. sp.1 of Prauss 2012(b), Bed G, MR97-2(A), M37/4, MPK 14701. Two overlapping
specimens. Note the presence of omphali and absence of a kalyptra;
6. Leberidocysta chlamydata (Cookson & Eisenack, 1962) Stover & Evitt, 1978, local Bed II, MR97-
21(A), J47/1, MPK 14702;
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7. Canninginopsis? lindseyensis sp. nov., Bed C, MR97-11(B), W33/2, MPK 14703. Note the
interpretation of precingular tabulation and sulcal notch (arrow);
8. ?D. globulus, Bed E, MR97-8(B), G27/2, MPK 14704. There are two precingular plates with
sutures on the specimen and an archaeopyle involving some detached dorsal precingular plates;
9. Bosedinia cf. sp.1 of Prauss 2012(b), Bed E, MR97-8.5(A), U39/4, MPK 14705. Note the
operculum, probably including apical and three intercalary plates;
10. Bosedinia cf. sp.1 of Prauss 2012(b), or an endocyst of Subtilisphaera spp., Bed G, MR97-2(C),
O39/4, MPK 14706;
11. Bosedinia laevigata (Jiabo, 1978, ex He & Qian, 1979) He, 1984, Bed E, MR97-8(A), R31/0,
MPK 14707;
12. B. laevigata, Bed C, MR97-11(E), H57/3, MPK 14708. Note the operculum, including apical and
probable intercalary 2a and 3a plates (indicated);
13. B. laevigata, Bed C, MR97-11.5(A), W40/1, MPK 14709;
14. Trithyrodinium maculatum sp. nov., Bed C, MR97-11.5(A), E45/0, MPK 14710. Endocyst. Note
the three anterior intercalary plates archaeopyle with at least two attached operculum plates, and the
endocyst wall marked by ring-shaped indentations;
15. T. maculatum, Bed C, MR97-10(A), B49/2, MPK 14711. Endocyst. Focus on probable intercalary
plates at top of specimen;
16. Fromea amphora Cookson & Eisenack, 1958, Bed C, MR97-11.5(A), N32/1, MPK 14712;
17. B. laevigata, Bed C, MR97-11.5(A), J43/3, MPK 14713;
18. T. maculatum, Bed F, MR97-3(A), S32/3, MPK 14662. Holotype. Note comparable ring-shaped
indentations on endophragm and periphragm. The likely position of the three intercalary plate
archaeopyle (operculum plates detached) is indicated;
19. T. maculatum, Bed B, MR97-12(D), K29/2, MPK 14714. Endocyst. Note the two plate intercalary
archaeopyle (2a [isodeltaform hexa-type] and 3a plates detached);
20. Sentusidinium ringnesiorum (Manum & Cookson, 1964) Wood et al., 2016, Bed C, MR97-
11.5(A), K42/2, MPK 14715;
21. B. laevigata, Bed E, MR97-8(B), X34/0, MPK 14716.
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Fig. 8. Kerogen (>10 μm fraction) photographs from Melton Ross. All fields of view are at 200 x 200
μm. Photographs 2, 11 and 12 are from stained kerogen slides.
1. Bed G, MR97-2(K), O42. Note the dominance of Amorphous Organic Matter (AOM);
2. Bed F, MR97-4(A), N42. Note the dominance of dinoflagellate cysts (d.c.) and small phytoclasts;
3. Bed F, MR97-5(K), N42. Note the presence of translucent (brown) phytoclasts and AOM;
4. Bed E, MR97-8(K), O42. Note the dominance of AOM;
5. Bed E, MR97-8.5(K), O42;
6. Bed D, MR97-9(K), O42. Note the prominence of dinoflagellate cysts, including Spiniferites
ramosus (S.r.);
7. Bed C, MR97-11(K), O42. Note the dominance of AOM;
8. Bed C, MR97-11.5(K), N42;
9. Bed B, MR97-12(A), O43. Note dinoflagellate cysts, including Oligosphaeridium totum (O.t.),
phytoclasts and pyritic mineral material;
10. Local Bed VII, MR97-15(A), O43;
11. Local Bed II, MR97-21(A), O42. Note dinoflagellate cysts, including Palaeohystrichophora
infusorioides (P.i.), Subtilisphaera spp. (Subt.) and Leberidocysta chlamydata (L.c.);
12. Local Bed I, MR97-22(A), O41. Note dinoflagellate cysts, including Litosphaeridium
siphoniphorum (L.s.).
Fig. 9. Van Krevelen-type diagram for the Melton Ross samples analysed for Rock-Eval pyrolysis.
Blue-filled circles are data for the > 1 wt.% TOC samples. Open circles are data for the < 1 wt.%
TOC samples; the latter are unreliable, due to probable elevation by occluded CO2 within carbonate of
inorganic origin, in addition to that derived from early diagenesis of organic matter.
Fig. 10. Lithological succession in the Bridge Creek Limestone Member of the Greenhorn Formation,
on the north side of the Pueblo Reservoir State Recreation area, west of Pueblo, Colorado, United
States. This section contains the Global boundary Stratotype Section and Point (GSSP) for the base of
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the Turonian Stage (38º 16‘ 56‖ N, 104º 43‘ 39‖ W) and the proposed GSSP for the base of the
Middle Turonian Substage (Bengtson et al. 1996; Kennedy et al. 2000, 2005; Dodsworth & Eldrett
2019). (a) metres above / depth below base of Bridge Creek Member; (b) chronostratigraphy; (c)
lithostratigraphy; (d) lithology; (e) δ13Corg, data from Bowman & Bralower (2005). Peaks ‗a‘, ‗b‘ and
‗c‘ in the OAE-2 excursion follow Jarvis et al. (2006); (f) summary of proposed regional correlation
with sections from NW Europe (this paper); (g) Pueblo ammonite zones after Kennedy & Cobban
(1991), Cobban (1993) and Kennedy et al. (2000). The Watinoceras devonense, Pseudaspidoceras
flexuosum and Vascoceras birchbyi units are often treated as subzones of a Watinoceras coloradoense
Zone or Watinoceras spp. Zone in published literature; (h) Pueblo inoceramid bivalve zonation by
Walaszczyk & Cobban in Kennedy et al. (2000); (i) Pueblo planktonic foraminiferal zones are from
Caron et al. (2005) and Keller & Pardo (2004); (j) subzones are from Keller & Pardo (2004). There is
a difference between Keller & Pardo (2004) and Caron et al. (2005) on the base of
Helvetoglobotruncana helvetica. This difference is marked by the vertical line in the figure; (k)
nannofossil zones are from Bralower (1988) and Bralower & Bergen (1998); (l), (m) dinoflagellate
cyst zones and subzones are from Dodsworth & Eldrett (2019). This diagram is adapted from
Dodsworth & Eldrett (2019). The main sources for NW Europe correlation are: Melton Ross, Wood &
Mortimore 1995, Wood et al. 1997, this paper, C.L. = Central Limestone; Misburg HPCF II quarry,
Hilbrecht 1986, Hilbrecht & Hoefs 1986, Prauss 2006, P.P. = pre-Plenus succession, P.B. = Plenus
Bed, triple = ‗triple band‘; Eastbourne, Jarvis et al. 2006, Pearce et al. 2009, Beds 1–8 compose the
Plenus Marls, the overlying Mead Marls occur within the Ballard Cliff Member and the Holywell
Marls occur within the Holywell Member.
Fig. 11. Scatter diagram plotting 13C -18O co-variance for the Melton Ross section to help elucidate
the primary versus diagenetic signals. Black circles = > 0.2 wt.% TOC; white circles = < 0.2 wt.%
TOC. Reference fields plotted from Moore (1989), with fields of well-preserved foraminifera (after
Veizer & Prokoph 2015; O‘Brien et al. 2017) most likely reflecting unaltered 13C -18O values
indicative of primary Late Cretaceous sea-water. Field of bulk rock 13C -18O values from CTB
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sections after Jarvis et al. (2011, 2015). Regression lines and R2 values plotted for all samples (black
line) and low TOC samples between samples MR97-11 and MR97-23 (dashed line).
Fig. 12. The stratigraphy of the Cenomanian–Turonian boundary interval in Norwegian Continental
Shelf, North Viking Graben exploration well 35/6-2 S (‗Grosso‘), drilled by StatoilHydro AS in 2009.
The proposed correlation to onshore outcrops is discussed in the main text. Shetland Group
lithostratigraphy of Gradstein & Waters (2016), dinoflagellate cyst zones and subzone of Olde et al.
(2015a) and Dodsworth & Eldrett (2019). The last occurrence (LO) of Carpodinium obliquicostatum
is taken here as a proxy for the top of the Adnatosphaeridium tutulosum Subzone. FO = First
Occurrence; NWG = NW Germany; GR = Gamma Ray, American Petroleum Institute (API) units.
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CLIP
CentralAtlantic
Western InteriorSeaway
30°N
60°N
0°NTrans-Saharan
Seaway
NeoTethys
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0 km 30
0 miles 20
Nor thSea UK
Speeton
Bridlington
Welton
South FerribyElsham
Caistor
Louth
Tetford
SouthThoresby
ClaxbyWelton le Marsh
HunstantonHeacham
Hillington
Marham
HULL
KING’S LYNN
NORWICH
LINCOLN
A
BA
B
Cenozoic
Chalk & Red Chalk
Gault Clay
Lower Cretaceous
Pre-Cretaceous
EastKnapton
Flixton
BishopWilton
MarketWeighton
47/10-1
Bigby MeltonRoss
WashThe
BGS Trunchborehole
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3
Deepexcavations,as numbered by Wood et al. (1997) Trench
Old Melton Rossquarries
Working areas in 1997 and location of deep excavationsand the trench, exposing the expanded CTB succession; the MR97 samples -1 to -13 were collected from Site 4 while samples -14 to -23 were collected from Site 1
Old Melton Rossquarries
B121
1
railway
0 125 250
metres
375 500
N
4
21
Figure 3
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A
B
C
E
F
D
G H
0
1
2
23
222120
19
18.5
18
17
16
15
14
13
12
13
2
45
6 -11.5
m Location: Melton Ross Quarry
Sample MR97-
BlackBand
VII
?VI
III
?III
?V-?IV
Lithostratigraphy Lithology
Figure 4
2 3
δ13C carbonate
a
?b
c
4
-6 -3 -4 -5
1 2 3
Biostrat.Age
Key: A. P. M. = Adrian’s Pair of MarlsF. C. F. = Ferriby Chalk Formation Bivalvia (inoceramid) debris Pyrite nodule Prominent macrobioturbation Prominent microbioturbation
(Chondrites)
Composite log of 1997 deep excavations, Site 1 (base of section to Central Limestone)and Site 4 (Bed B to top of section)
Chondrites II
Chondrites I
δ18O carbonate
5
-7
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9 11
50μm
Figure 5
7
21 3
65
4
10
18
12 14 15
17 19 20 21
8
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50μm
Figure 6
9
21
4
10
765
11
12 13 14
8
3
1IlI
1II
1IIII
2IlI
1c2c
lp
3c4c
5c6c
3IlI4IlI
2II4II5II ?6II
5IlI?6IlI5IlI
?s
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11
Figure 7
7
21 3
65
4
10
19
12
16 17
18 20 21
13
15
50μm
14
9
8
1a 3a2a2a
3a
2a
3a
2a
1a2a
3a
1a 3a
1IIII
3II2II4II
5II6II1II
2a 3a
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Figure 8
8
21 3
654
10 11 12
97
O.t.
pyrite
L.s.
L.c.
P.i.
Subt.
pyritic material
brownphytoclast
blackphytoclast
AmorphousOrganic Matter
S.r.
brownphytoclast
d.c.
AOM
blackphytoclast
MR97-2Bed G
MR97-4Bed F
MR97-5Bed F
MR97-8, Bed E MR97-8.5Bed E
MR97-9Bed D
MR97-12Bed B
MR97-11.5Bed C
MR97-11Bed C
MR97-15Bed VII
MR97-21Bed II
MR97-22Bed I
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0
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0 10 20 30 40 50 60 70 80 90 100
HY
DR
OG
EN
IND
EX
(H
I, m
g H
C/g
TO
C)
OXYGEN INDEX (OI, mg CO2/g TOC)
TYPE I
TYPE II
TYPE IV
TYPE III
l
II
III
IV
Geochemical Services Group, 143 Vision Park Blvd., Shenandoah, Texas 77384 • Phone: 281-681-2200 • Fax: 281-681-0326 • Email: [email protected]
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125
122120118116113
109
105
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97
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12
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-28 -26 -24 -22
δ13Corg
a. c.b. d. e.
AmmoniteZones
f.
Planktonicforaminifera
Zones
h.
Calcareousnannoplankton
Zones
j.
Nigericas scotti(inferred)
P. flexuosum
V. birchbyi
Mytiloidesmytiloides
Mytiloideskossmati
Mytiloides puebloensis
Mytiloides hattini
Inoceramuspictus
InoceramidZones
g.
Adnatosphaerid. tutulosum
Isabelidinium magnum
Senoniasphaera turonica
Cyclonephelium membraniphorum
Dinoflagellatecyst
Zones
k.
Dinoflagellatecyst
Subzones
l.
LitosphaeridiumsiphoniphorumRotalipora
cushmani
A. tutulosum
Dicarinellahagni
Heterohelixmoremani
G. bentonensisRotaliporaExtinction
?
Mytiloidessubhercynicus
PlanktonicforaminiferaSubzones
i.
Axopodorhab.albianus
Rhagodiscusasper
Eprolithus florialis
Collignoniceraswoollgari
Mammitesnodosoides
Neocardiocerasjuddii
Sciponocerasgracile
Metoicocerasmosbyense
Watinocerasdevonense
Helvetoglobotrunc.helvetica
Whiteinella archaeocretac.
NW Europecorrelation
m.
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b
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Figure 10
C.L.
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35/6-2 SField: Grosso
Location: Norwegian CSOperator: StatoilHydro AS
Scale: 1:750
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Biostratigraphic Comments
LO Carpodinium obliquicostatum
FO common Cyclonephelium compactum-membraniphorum. Eurydinium saxoniense & Alterbidinium daveyi
LO common / consistent Litosphaeridium siphoniphorum, LO Pterodinium crassimuratum
LO Pterodinium cingulatum granulatum
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2616 D/C
2625 D/C
2646 D/C
2655 D/C
2664 D/C
2673 D/C
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Fish Shale (NWG) /Beds C - H
(eastern England)
Bed B
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nStratigraphic Summary Chart
Figure 12
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Locality: Melton Ross. Mortimore Hildreth Wood et Dodsworth Wood & Jeans et al. Jeans et al. Hart et al. JefferiesLithostratigraphy and sampling. This paper 2014 1999 al . 1997 1996 Mortimore 1991 1991 1991 & 1963(Not to scale.) 1995 1993
Melton Ross S. Ferriby Bigby M. Ross S.F./Flixton M.R./S.F. Flixton S. Ferriby S. Ferriby S. FerribyMember Lithology Sample Bed Sub‐unit
limestoneMR97‐1 Bed H gungy marl Unit 8 Bed 10 Bed H Bed 6 Bed 8 Bed vMR97‐2 Bed G Bed 9 Bed G ?15 to ?17 Bed 7
light MR97‐3 Bedsmarl MR97‐4 Bed F Unit 7 Bed 8 Bed F Bed 5 11 to ?14 Bed 6 Bed iv Bed iv
Flixton MR97‐5Member MR97‐6(Black dark MR97‐7 Bed E Bed 7c Bed E Bed 10
Band Sub‐ marl MR97‐8
member) MR97‐8.5 BlackMR97‐9 Bed D Band Unit 6 Bed 7b Bed D Bed 4 Bed 9 Bed 5 Bed iii Bed iiiMR97‐10 C3 C3 Bed 8MR97‐10.5 Bed C C2 Bed 7a Bed C C2 Bed 7
dark MR97‐11 C1 C1 Bed 6marl MR97‐11.5
MR97‐12 B4 Unit 5 Bed 6b (u) Bed 3 Bed 5light silty Bed 6a
Flixton marl MR97‐13 Bed B B3 khaki Unit 4 Bed 5b Bed B Bed 2 Bed 4 Bed 4 Bed ii Bed iiMember B2 marl Unit 3 Bed 5a (l)(Plenus B1 Bed 4
Marls Sub‐ limestone MR97‐14 A3 (CL) Unit 2 (CL) Bed 3 (CL) lst limestone Bed 3 (lst)member) Bed A A2 Unit 1 Bed 2 Bed A Bed 1 Bed 3 Bed 2 Bed i Bed i
A1 Bed 1 Bed 2MR97‐15 to 22 Beds I‐VII Beds I‐VII A. S.
Louth Mbr. limestone MR97‐23 Ferriby Chalk Formation Bed 1 Bed 1
Key: A.S. = 'Anomalous Succession'; CL, = 'Central Limestone'; lst = limestone (chalk); l = lower; u = upper. = Stratigraphic breakTable 1. Lowermost part of the Welton Chalk Formation: A comparison of bed nomenclature.
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Locality: Melton Ross.(Not to scale.) Kerogen Terrigenous palynomorphs Marine palynomorphsLithostrat. Lithology Sample Counts per TOC AOM P'clasts Palyn. Gymnosp. Pterido. Angiosp. T/M P/G Dinoflag. Prasinoph. Acantho. Dinoflag.
g (total) % % % % % % % ratio ratio % % % diversitylimestone
Bed H MR97‐1 34,320 0.21 71 12 17 9.7 0.3 < 0.3 0.1 0.24 81.7 5.7 2.7 63Bed G MR97‐2 89,760 3.47 92 4 4 11 2.3 0.3 0.137 0.34 68.3 6.3 11.7 62
light MR97‐3 6,688 0.18 58 13 29 3 < 0.3 < 0.3 0.03 0.21 95.3 0.7 0.7 69Bed F marl MR97‐4 1,470 0.12 9 49 42 < 0.3 < 0.3 0 0 0.31 98.7 1.3 0 53
MR97‐5 8,668 0.34 41 33 26 2.7 < 0.3 0 0.027 0.16 96.3 0.7 0.3 67MR97‐6 17,204 0.4 42 30 28 5 1 0.3 0.063 0.25 91.3 1.3 0.7 79
Bed E dark MR97‐7 30,360 0.67 31 33 36 9.7 0.3 < 0.3 0.1 0.36 82.7 4.7 2.3 68marl MR97‐8 85,184 2.18 83 8 9 9.7 2.6 < 0.3 0.123 0.32 77.3 6 4 71
MR97‐8.5 219,648 2.74 71 11 18 12.3 0.3 < 0.3 0.127 0.35 82.7 1.7 2.7 72Bed D MR97‐9 23,518 0.36 19 30 51 < 0.3 0 0 0 0.03 100 0 0 63
MR97‐10 123,024 2.19 81 5 10 10 0.3 0.3 0.107 0.29 86.7 1 1.7 72Bed C MR97‐10.5 53,284 0.7 27 29 44 0.3 0 < 0.3 0.003 0.04 99 0 0.3 65
dark MR97‐11 101,552 1.43 63 16 21 4 < 0.3 0.7 0.047 0.31 92.3 0.7 2.3 79marl MR97‐11.5 87,296 2.2 71 15 14 7.3 < 0.3 0.3 0.077 0.35 90.7 0.7 1 76
Bed B MR97‐12 288 0.2 0.3 0 0 0.003 0.01 99.7 0 0 33MR97‐13 28 0.12 1.4 0.4 0 0.018 0.06 96.5 0 0.7 33
Bed A limestone MR97‐14 <1 0.1 No reliable kerogen 2MR97‐15 0 0.15 counts from sample ‐12 0
Bed VII MR97‐16 <1 0.11 to sample ‐19, due to 1light MR97‐17 27 0.14 sparse recovery and 0 0 0 0 0 100 0 0 4marl MR97‐18 5 0.13 pyritic material. 0 0 0 0 0 100 0 0 1
Bed ?VI MR97‐18.5 8 0.21 0 0 0 0 0 100 0 0 2Bed ?IV‐V MR97‐19 88 0.14 0 0 0 0 0 100 0 0 1Bed ?III MR97‐20 95 0.16 1 89 10 0.3 0 0 0.003 0.72 99.3 0.3 0 32Bed II MR97‐21 12,408 0.28 2 26 72 0.7 < 0.3 < 0.3 0.007 0.45 98.3 1 0 77Bed I MR97‐22 53 0.12 4 90 6 1 0 0 0.01 0.66 97.3 1.7 0 33
limestone MR97‐23 N/A 0.04Table 2. Summary of palynomorph groups, > 10 micron kerogen counts and total organic carbon (TOC). Shaded rows highlight samples with > 1 % TOC.
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Locality: Melton Ross.(Not to scale.) Gonyaulacoid dinoflagellate cystsLithostrat. Lithology Sample L. siphoniphorum C. compactum‐memb. Pterodinium spp. Spiniferites spp. O. totum Dinoflagell.? type D
% c.p.g. % c.p.g. % c.p.g. % c.p.g. % c.p.g. % c.p.g.limestone
Bed H MR97‐1 0 0 42.7 14655 0.7 240 6 2059 0 0 1.3 458Bed G MR97‐2 0 0 11.7 10502 2.3 2064 9.3 8348 0 0 0.7 598
light MR97‐3 0 0 13 869 10 669 28.3 1893 < 0.3 P 2 134Bed F marl MR97‐4 0 0 19.7 290 3.7 54 18.7 274 < 0.3 P 9 132
MR97‐5 0 0 21.7 1881 4 347 27.7 2401 0.7 58 4 347MR97‐6 0 0 18 3097 3.7 637 22.3 3836 < 0.3 P 1.7 287
Bed E dark MR97‐7 0 0 15.3 4645 3.7 1123 14.7 4463 0 0 1 304marl MR97‐8 0 0 15.7 13374 4 3407 16.7 14226 < 0.3 P 0.3 284
MR97‐8.5 0 0 23.3 51178 2.7 5930 9.3 20427 0 0 0.3 732Bed D MR97‐9 0 0 35 8231 3.3 776 33 7761 1.3 306 1.3 314
MR97‐10 0 0 23.7 29157 3 3691 12 14763 < 0.3 P 0.3 410Bed C MR97‐10.5 0 0 32.0 17050 4.7 2504 25.7 13694 0.3 158 0.7 355
dark MR97‐11 0 0 16 16248 4 4062 25.4 25794 0.7 656 0.7 677marl MR97‐11.5 0 0 13 11348 5 4365 17.3 15102 3.3 2612 1 873
Bed B MR97‐12 ? < 0.3 ?P 13.7 39 < 0.3 P < 0.3 1 40.3 116 16.3 47MR97‐13 0 0 29.3 8 0 0 23.3 7 5 1 1.1 <1
Bed A limestone MR97‐14 0 0 0 0 0 0 0 0 0 0 0 0MR97‐15 0 0 0 0 0 0 0 0 0 0 0 0
Bed VII MR97‐16 0 0 0 0 0 0 0 0 0 0 50 <1light MR97‐17 0 0 0 0 0 0 0 0 0.4 1 98.1 26marl MR97‐18 0 0 0 0 0 0 0 0 0 0 100 5
Bed ?VI MR97‐18.5 0 0 0 0 0 0 0 0 0 0 94.7 8Bed ?IV‐V MR97‐19 0 0 0 0 0 0 0 0 0 0 100 88Bed ?III MR97‐20 1.7 2 0.3 1 1 1 5.4 5 0 0 48 46Bed II MR97‐21 2.7 335 < 0.3 P 4.7 583 29 3598 0 0 1 124Bed I MR97‐22 1 1 0.3 1 4.3 2 13.3 7 0 0 3 2
limestone MR97‐23Table 3. Relative and absolute abundance of selected gonyaulacoid dinoflagellate cysts (G‐cysts), and Dinoflagellate? type D of Ioannides (1986).ACCEPTED M
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Locality: Melton Ross.(Not to scale.) Peridinioid dinoflagellate cystsLithostrat. Lithology Sample P. infusorioides S. pontis‐mariae E. saxoniense Bosedinia cf. sp. 1 T. suspectum Ginginodinium? sp.A
% c.p.g. % c.p.g. % c.p.g. % c.p.g. % c.p.g. % c.p.g.limestone
Bed H MR97‐1 < 0.3 P 5 1716 4.7 1613 1 343 7 2402 1.7 583Bed G MR97‐2 0.3 269 4 3590 8 7180 5.7 5116 3 2693 1 898
light MR97‐3 11.3 756 3.3 221 1.7 114 0.3 20 2 134 0.7 47Bed F marl MR97‐4 9.3 137 5.7 84 1.7 25 0.3 4 4 59 0.3 4
MR97‐5 7.7 667 2.7 234 1.7 147 2 173 0.3 26 0 0MR97‐6 9 1548 5.3 912 2.7 465 2 344 2 344 0.7 120
Bed E dark MR97‐7 3.3 1002 8.7 2641 5.3 1609 5 1518 4.7 1427 1.7 516marl MR97‐8 3.7 3152 10.7 9115 2.3 1959 3 2556 2.7 2300 1 852
MR97‐8.5 1.7 3734 18.7 41074 2 4393 2.3 5052 2.3 5052 1.3 2855Bed D MR97‐9 1 235 0.7 165 0.3 71 0 0 0.7 165 0 0
MR97‐10 1 1230 14.7 18085 1 1230 6 7381 1.7 2091 < 0.3 PBed C MR97‐10.5 1.3 693 < 0.3 P 0.3 160 0 0 1.7 906 0 0
dark MR97‐11 5 5078 16.7 16959 0.7 711 1.3 1320 4 4062 0 0marl MR97‐11.5 3 2619 18.7 16324 1 872 2 1746 5 4365 0.3 262
Bed B MR97‐12 0 0 < 0.3 P 0 0 0 0 1 3 0 0MR97‐13 3.2 9 0.7 1 0 0 0.4 1 1.1 1 0 0
Bed A limestone MR97‐14 0 0 0 0 0 0 0 0 0 0 0 0MR97‐15 0 0 0 0 0 0 0 0 0 0 0 0
Bed VII MR97‐16 0 0 0 0 0 0 0 0 0 0 0 0light MR97‐17 0 0 0 0 0 0 0 0 1.1 1 0 0marl MR97‐18 0 0 0 0 0 0 0 0 0 0 0 0
Bed ?VI MR97‐18.5 0 0 0 0 0 0 0 0 0 0 0 0Bed ?IV‐V MR97‐19 0 0 0 0 0 0 0 0 0 0 0 0Bed ?III MR97‐20 13.3 13 17.7 17 0 0 0 0 1.3 1 0 0Bed II MR97‐21 32 3971 10 1241 0 0 < 0.3 P 0.7 87 < 0.3 PBed I MR97‐22 13 7 41.3 22 ?1 ?1 0 0 2.3 1 0.3 1
limestone MR97‐23Table 4. Relative and absolute abundance of common/abundant peridinioid dinoflagellate cysts (P‐cysts).ACCEPTED M
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