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changes in the Permian (Lopingian) Zechstein deposits of northeast England. Review of
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The use of spore-pollen assemblages to reconstruct vegetation changes in the Permian(Lopingian) Zechstein deposits of northeast England

3

Martha E. Gibsona* and Charles H. Wellmana

45aDepartment of Animal & Plant Sciences, University of Sheffield, Alfred Denny Building,Western Bank, Sheffield S10 2TN, UK.

6*marthae.gibson@gmail.com

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8

ABSTRACT

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New boreholes have enabled, for the first time, extensive palynological sampling through theentire Lopingian Zechstein sequence of northeast England. Palynomorph assemblages havebeen recovered from throughout the sequence from all five of the evaporation-replenishment
cycles (EZ1-EZ5). These assemblages are dominated by pollen grains, with rare trilete spores,
and even rarer marine forms such as acritarchs and foraminiferal test linings. The assemblageof pollen grains is of low diversity (35 species) and dominated by taeniate and non-taeniate
bisaccate pollen. The assemblage varies to only a limited extent both within and between
cycles, although some minor variations and trends are documented. Based on the compositionof the dispersed spore-pollen assemblages, and previous work on the Zechstein megaflora, thehinterland vegetation is interpreted as being dominated by conifers that inhabited a semi-arid
to arid landscape. This flora is shown to have persisted throughout the entire Zechsteinsequence, despite previous assertions that it disappeared as conditions became increasingly
drier over the course of the latest Permian.

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Key words: palaeobotany, palynology, pollen, spores, vegetation change, Permian.

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1. Introduction

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The Lopingian Zechstein Sea was a large intercontinental sea located in Pangaea justnorth of the equatorial Central Pangaean Mountain Range. A remarkable stratigraphicalsequence deposited in the sea is dominated by carbonate/evaporite cycles. The sequence canbe divided into seven cycles at the centre of the basin, although this is reduced to five cycles
towards the margins (EZ1-5) (Figure 1). These cycles essentially represent a sequence of largescale evaporation-replenishment events. The biota of the Zechstein Sea and surrounding
hinterlands is also remarkable. Marine biotas are impoverished due to high salinities. Although
reefs developed during the early cycles, these diminished and then vanished as conditions
became more saline and harsher during the later cycles. Palaeobotanical and palynologicalstudies reveal that a typical Euramerican flora occupied the hinterlands, low lying areas andriparian environments around the Zechstein Sea during the early cycles (EZ1-2). However,
hitherto there has been little floral evidence in later cycles (EZ3-5) and it is often assumed thatvegetation diminished as harsher desert conditions prevailed as the climate became warmer
and drier. It is of particular interest that the sequence accumulated during the build up to theend-Permian mass extinction (Erwin, 1993; Benton and Twitchett, 2003; Erwin, 2006; Hallam
and Wignall, 1997) and there is some evidence that the event may itself be reflected in thehighest cycle toward the centre of the basin (e.g. García-Veigas et al., 2011).

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Recently, new boreholes have become available that penetrate the younger cycles (EZ35) at the margins of the Zechstein Sea in northeast England. These have yielded, for the firsttime, rich palynomorph assemblages from these younger cycles. This paper reports on aquantitative analysis of these palynomorph assemblages, in addition to those from older cycles(EZ1-2) collected at outcrop. The new data provides evidence for the nature of the hinterland
vegetation from throughout all five cycles developed at the margins of the basin. This has
enabled reconstruction of the vegetation changes that accompanied the evaporationreplenishment cycles observed in the sedimentary sequences.
Cycle- 25 1 Ma

Durham Sub.basin

Yorkshire Sub.basin

Brockelschiefer

Brockelschiefer

Roxby Formation

Littlebeck Anhydrite Formation

Sequence

Roxby Formation

EZ5

ZS7

Sleights Siltstone
Sneaton (Halite) Formation
EZ4

Sherburn (Anhydride) Formation

Upgang Formation

c.

::,

Carnallitic MarlC)C:

Boulby Halite

Boulby Halite

Billingham Anhydrite Formation

Billingham Anhydrite Formation

e

C:

C:
.!l! .!l!
Cl

E
...
(I)

0.

C:

'ii !!
0...J

Sherburn (Anhydrite) Formation

EZ3

Cl)

~

Seaham Formation

0
Cl>

N

Edlington Formation
EZ2
Roker Formation--------------------------------------Concretionary Limestone Member

Hartlepool Anhydrite Formation
EZ1-258 Ma

Ford Formation
Raisby Formation
Marl Slate Formation

ZS6

Carnallitic Marl
ZS5Brotherton Formation
Grauer Salzton Formation

Fordon Evaporite Formation

ZS4

Edlington Formation
Kirkham Abbey Formation

ZS3

Hayton Anhydrite
Cadeby Formation____ Sprotbrough Member __Wetherby Member

Marl Slate Formation

ZS2
ZS1

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Figure 1. Stratigraphy and correlation of the UK Zechstein deposits between the Durham Subbasin and Yorkshire Sub-basin. Approximate Zechstein Group dates taken from Menning et al.
(2005, 2006).

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2. Geological Setting

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In western and central Europe, a Lopingian to early Triassic marine-continental
transition accumulated in the intracontinental Southern Permian Basin (SPB). The SPB evolvedon the former Variscan foreland in the latest Carboniferous-early Permian (Ziegler, 1990). The
depositional area has a width of ca. 600 km and trends ca. 2,500 km WNW-ESE from northeast
England, stretching across the southern North Sea, and into northern Germany and Poland. The
SPB is delineated by several Variscan Highs including the London–Brabant, Rhenish andBohemian massifs in the south, and the Mid North Sea, and Ringkøbing–Fyn highs in the north(Ziegler, 1990; Geluk, 2005; Peryt et al., 2010).

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Within the SPB tectonism initially had a relatively minor influence on the depositionof the Lopingian-early Triassic deposits. Rather, evolution of the SPB is considered to havebeen influenced by thermal relaxation of the lithosphere (van Wees et al., 2000) causing
differential subsidence across the basin. However, the role of tectonism increased inimportance in the latest Permian-earliest Triassic (uppermost Zechstein-Buntsandstein) when
tectonic pulses began to affect sedimentary successions.

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The initial transgression of the Zechstein Sea was a consequence of rifting in the ArcticNorth Atlantic, accompanied by a contemporaneous rise in global sea levels. This resulted inflooding of the SPB via the Viking Graben System (Ziegler, 1990). Although this flooding
pathway had existed previously and was responsible for minor and short-lived marine
incursions during the middle Permian Upper Rotliegend (Legler and Schneider, 2008), theinitial Zechstein transgression was a much more catastrophic flooding event. It is represented
by the deposition of the ‘Kupferschiefer’ (copper shale), or Marl Slate in the UK, whichindicates permanent flooding of the sub-sea-level SPB (Glennie and Buller, 1983; van Wees etal., 2000).

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The Kupferschiefer/Marl Slate was deposited under basin-wide euxinic conditions(Pancost et al., 2002; Paul, 2006) and is a 0.5 m thick layer of black shale across the basin thatprovides an excellent marker horizon utilised in regional correlation (Geluk, 2005; Doornenbal
and Stevenson, 2010). A Re-Os age of 257.3 ± 1.6 Ma for the Kupferschiefer in central
Germany (Brauns et al., 2003) indicates a Lopingian (early Wuchiapingian) age for the basal
Zechstein. However, this Re-Os age has been challenged (Słowakiewicz et al., 2009). Datingof the Zechstein in central Germany places the Zechstein Transgression at 258 Ma (Menning
et al., 2005, 2006) (Figure 1).

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During the Lopingian the SPB was located within the supercontinent Pangaea in thenorthern hemisphere desert belt at 10-30°N (Glennie, 1983). Zechstein deposition took place
under arid and semi-arid climate conditions, the influence of which is documented by up toseven cycles of sedimentation and stacked evaporation cycles (Figure 1). These provide aframework for lithostratigraphical subdivision and facilitate basin-wide correlation (RichterBernburg, 1955). While the Zechstein Sea existed, the SPB was filled with approximately
2000m of siliciclastics, carbonates and evaporites (Peryt et al., 2010) following the classic
model of cyclical chemical precipitation in a giant saline basin of Richter-Bernburg (1955). At
its maximum extent, towards the centre of the Zechstein Sea, the Zechstein Group consists ofseven evaporation cycles, the Werra (Z1), Staßfurt (Z2), Leine (Z3), Aller (Z4), Ohre (Z5),
Friesland (Z6) and Fulda (Z7) cycles.

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The entire Zechstein sequence is considered to be of Lopingian age and accumulated
within a period of 2.8-3.5 Myr (Szurlies, 2013; Menning et al., 2005), 5 Myr (Menning et al.,2006), or up to 9 Myr (Słowakiewicz et al., 2009) (Figure 1). There is no consensus yet for one
timescale for all of the Zechstein sequence and it is likely that different regions of the basin
were deposited at different times. The British Zechstein is yet to be dated and, so far, attempts
to match the Zechstein in the UK to the global chronostratigraphic timescale have been
unsuccessful (D. Grocke pers. comm.). Consequently, the dates presented here are based on
the latest magnetostratigraphical dating from the German/Dutch Zechstein successions (i.e.
Szurlies, 2013) and carbon isotope studies (e.g. Słowakiewicz et al., 2009).

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The UK Zechstein crops out over northeast England from Country Durham south toNottinghamshire (Figure 2). These deposits accumulated in two sub-basins, separated by the
Cleveland High, which ran along the western margin of the sea: the Yorkshire Sub-basin to thesouth and the Durham Sub-basin to the north (Smith, 1989). The sequence is divided into fiveevaporation cycles represented by EZ1-5 (Figure 1). The Zechstein Group is separated fromthe deposits of the Bakevellia Sea to the west by the low-lying, gently undulating topographyof the Protopennines (Stone et al., 2010).

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The Zechstein displays an overall regressive and hypersaline trend, with the thicknessof strata thinning towards the margins of the SPB. As such, only the first three cycles display
a full cycle of lithologies including marine clastics, carbonates and evaporites. The upper cycles

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are dominated by clay and siltstones and generally contain fewer evaporites. These evaporites
formed in more localised depressions and do not extend basin-wide (Smith, 1989).

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The Kupferschiefer (Marl Slate) represents one of three regionally extensive marker
horizons used in correlation of the Zechstein (Geluk, 2005), the others being the ‘Grauer
Salzton’ (Grey Salt Clay/Illitic Shale) at the base of Z3, and the ‘Roter Salzton’ (Red Salt
Clay/Carnallitic Marl Formation) at the base of Z4. All three markers are assumed to represent
nearly synchronous flooding events (Doornenbal and Stevenson, 2010).

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Zechstein lithostratigraphy is well-defined and correlated throughout the interior of the
SPB for cycles Z1 to Z3. However, correlation of the upper Zechstein is more difficult withdisagreement regarding correlation of the Z4 of Poland (PZ4a-e) with Z5-lower Z7 in Germany(Wagner and Peryt, 1997; Käding, 2000; Peryt et al., 2010). In Germany, the ZechsteinBuntsandstein boundary is placed at the base of the first prominent sandstone directly above
the Z7 Formation (Röhling, 1993 and references therein).

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Zechstein deposition terminated when the connection with the Boreal Ocean wasinterrupted by major clastic influx into the SPB (Ziegler, 1990). The SPB initially transitioned
into an extensive sabkha environment with isolated salinas, later becoming an extensive inland
playa lake in which the Lower Buntsandstein Subgroup was deposited (Hug, 2004; Hug and
Gaupp, 2006). Salt precipitation progressively retreated further into the basin centre during theuppermost Zechstein. Tectonism resulted in downwarping of the central parts of the basin withuplift creating minor unconformities at its margins (Geluk, 1999). Zechstein sedimentation
finally ended with the progradation of the uppermost part of EZ5/Z7, a sequence of fine-grained
clastic facies formerly named the ‘Bröckelschiefer’ (Peryt et al., 2010).

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Figure 1 illustrates the general stratigraphical sequence of the Zechstein of northeast
England. Figure 2 illustrates the location of the Zechstein sequences analysed in this study and
Figure 3 illustrates the stratigraphy and correlation of the Zechstein sequences. Fulllithostratigraphical and facies descriptions are provided in Appendix A of the online
supplementary information. It is important to note that in northeast England only Cycles 1-3
are exposed. Later cycles (Cycles 4-5) and the Permian-Triassic boundary can only be accessedby boreholes and are comparatively understudied.

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3. Previous palaeobotanical and palynological studies

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The palaeobotany of the British Zechstein sequence is well documented and comprises
a typical Euramerican flora dominated by gymnosperms (Stoneley, 1958; Schweitzer, 1986;
Cleal and Thomas, 1995). Rich palynomorph assemblages have previously been describedfrom the Zechstein of northeastern England (Wall and Downie, 1963; Clarke, 1965) as well as
equivalent deposits from the Bakevillia Sea of northwest England (Clarke, 1965; Warrington,2008) and the Kings Court outlier in Ireland (Visscher, 1971, 1972). The palynology of other
Permian strata in the UK has been reviewed by Warrington and Scrivener (1988) and
Warrington (2005). All of these studies report mainly pollen assemblages (with subsidiary
spores) but include rare evidence for marine elements such as acritarchs and foraminiferal test
linings.

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Wall and Downie (1963) reported the presence of marine acritarchs in Zechstein
deposits from Ashfield Brick Pit, Conisborough, Yorkshire. They were rare, with theassemblages dominated by pollen, and included relatively simple forms such as Veryhachiumand Micrystridium.

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The most comprehensive review of British Zechstein palynology is Clarke (1965).
Spore-pollen assemblages from the Hilton Plant Beds, Kimberley Railway Cutting andHaughton Hall Boring in Nottinghamshire were reviewed. Thirty-three taxa were reported,with assemblages containing an overwhelming dominance of Lueckisporites virkkiae, with
Taeniaesporites noviaulensis and T. labdacus also being very abundant. Clarke reported thatthe most common non-taeniate taxa were Klausipollenites schaubergeri, Falcisporites zapfei,
Labiisporites granulatus and Illinites delasaucei. Monosaccate pollen were represented by
Perisaccus granulosus, P. laciniatus, Potonieisporites novicus, Vestigisporites minutus andNuskoisporites dulhuntyi, although none of these species are common. Monosulcates are only
represented by the genus Cycadopites. Clarke suggested that the similarity seen between
Zechstein assemblages and other co-eval assemblages from the UK indicated that the
Lopingian vegetation was fairly uniform throughout this region. Due to the palynologicalassemblages being derived from a very limited stratigraphical range it was not possible for
Clarke to comment on any temporal variation. Clarke also erected three variants of L. virkkiae
based on gross morphological differences. According to Clarke (1965) Variant A is the largest
and is described as having well-developed, distinctly separate proximal thickenings and welldeveloped sacci, being most similar to the holotype and L. microgranulatus Klaus 1963.
Variant B has sacci that are less well-developed and a small saccus offlap, being most similar
to L. parvus Klaus 1963. Variant C has a weakly developed proximal cap that is not completely
separated into two halves and is generally smaller with a more elongate corpus, being mostsimilar to L. microgranulatus “kleinere variante” Klaus 1963.

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Visscher (1971) reviewed the Permian and Triassic palynology of the Kingscourt
Outlier, Ireland, which is a correlative of the Bakevellia and Zechstein seas. The oldest
assemblage described (Assemblage 1) was found to closely correlate with Lopingian Zechstein
assemblages from western Europe. This assemblage is dominated by Lueckisporites virkkiae,
Jugasporites delasaucei, Klausipollenites schaubergeri and Limitisporites moerensis.
Perisaccus granulatus and Striatisaccus sp. are minor components. However, typical Zechstein
forms Falcisporites zapfei (Potonié and Klaus, 1954), Labiisporites granulatus andStriatopodocarpites richteri were notably absent, presumably due to the semi-isolated natureof deposits of the Kingscourt Outlier to the west of the Zechstein Basin. Visscher proposed azonation, based on ‘palynodemes’ of L. virkkiae, which represents a rare attempt to apply apalynology-based biostratigraphy to the Zechstein Group.

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More recent work on the Permian palynology of the UK has tended to focus onequivalents of the Zechstein deposits (Warrington and Scrivener, 1988; Legler et al., 2005;
Legler and Schneider, 2008; Warrington, 2005, 2008). For example, the Lopingian of Devon
yields poorly preserved bisaccate gymnosperm pollen (Warrington and Scrivener, 1988).
Lueckisporites virkkiae, Perisaccus granulosus, Klausipollenites schaubergeri, Jugasporites
delasaucei, Protohaploxypinus microcorpus and Lunatisporites spp. are found and are allcompatible with a Lopingian age. The Hilton Borehole, in the Valley of Eden in Cumbria,
yields the longest continuous section of Permian rocks in Britain and includes strata equivalent
to the Zechstein succession (EZ1) of eastern England (Jackson and Johnson, 1996). Warrington(2008) described palynomorph assemblages, dominated by pollen, but also containing algal
remains. Crucisaccates cf. variosulcatus and possibly Proprisporites pococki were reported
for the first time from the Upper Permian of the UK.

50°

Scarborough

e. 0
II

Shefild■

40 °' Nottingham

■
0

10

20

30

~

50km

40

40'

50°

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Figure 2. Outcrop map of the UK Zechstein deposits showing the location of the boreholes andoutcrop exposures considered in this paper. 1) Marsden Bay, 2) Claxheugh Rock, 3) Crime
Rigg Quarry, 4) Little Scar, 5) SM7 Mortar Hall, 6) SM11 Dove’s Nest, 7) SM4 Gough, 8)
SM14b Woodsmith Mine, 9) Pot Riding, 10) Levitt Hagg Hole, 11) Sandal House, 12) Ashfield
Brick Pit, 13) Salterford Farm, 14) Woolsthorpe Bridge, 15) Kimberley. Permian outcrops areshaded in grey.

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4. Materials and Methods

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The Zechstein sequence of northeastern England was extensively collected from both
outcrop and boreholes to gather a set of samples for palynological analysis from throughout
the sequence. A total of 192 samples were collected from six boreholes (SM4 Gough, SM7
Mortar Hall, SM11 Dove’s Nest, SM14b Woodsmith Mine North Shaft, Salterford Farm,
Woolsthorpe Bridge) and outcrop samples from the Yorkshire Sub-basin (Kimberley Railway
Cutting, Levitt Hagg Hole and Little Scar beach) and from the Durham Sub-basin (ClaxheughRock, Crime Rigg Quarry and Marsden Bay). Figure 1 shows the location of the boreholes andoutcrops discussed in this study. See Appendix B of the online supplementary information for

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a list of the borehole samples and their lithology, and Appendix C for a list of the outcrop
samples. Appendix D provides more details of the stratigraphic range of the sampled
localities/boreholes. The sampling resolution was determined by the frequency of lithologies
suitable for palynomorph preservation and by the extent of exposed sections/borehole material
available for sampling.

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The samples were palynologically processed using different techniques depending onwhether the lithology was clastic-carbonate or evaporite. For clastic-carbonate lithologies 20gof sample was prepared using standard palynological HCl-HF-HCl acid maceration techniques.
The residue was first top sieved at 10µm to detect the presence of acritarchs, then at 20µm, andsubjected to heavy liquid separation using zinc chloride to extract and concentrate the organic
residues. Evaporite lithologies were processed according to the method outlined in Gibson andBodman (2020), whereby samples are dissolved in hot water, boiled in concentrated 40% HF
for 15 minutes to remove any remaining clastics, then subjected to heavy liquid separation
using zinc chloride to extract and further concentrate any organic residue.

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For productive samples, the residues varied considerably in palynomorph
colour/opacity. In most samples the palynomorphs were beautifully preserved and of very low
thermal maturity (translucent yellow-orange). In some samples the palynomorphs were of very
low thermal maturity and appeared hyaline (e.g. Marl Slate of the Durham Sub-basin). These
were stained using Bismark Brown to improve contrast and visibility of palynomorphs. On theother hand, samples associated with evaporites were often opaque dark brown-black(frequently 98-100% PDI) (Goodhue and Clayton, 2010; Clayton et al., 2017). This darkening
is unlikely to be due to high thermal maturity as the Zechstein deposits have not been subjected
to deep burial and high heat flow and palynomorph assemblages above and below are of very
low thermal maturity (see Gibson and Bodman (2020)). It appears to result from a diagenetic
effect of preservation within evaporites. Where necessary, samples were oxidized using
Schulze’s reagent for up to 26 hours which cleared the palynomorphs to translucent orange.
Samples containing large quantities of amorphous organic matter (AOM) were treated withpulsed ultrasound treatment to break up the AOM and prevent it from obscuring palynomorphs
during analysis. The organic residue was strew mounted onto glass slides for light-microscopic(LM) analysis.

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Specimens were analysed and photographed using a QImaging (Model No. 01-MP3.3RTV-R-CLR-10) camera mounted on an Olympus BH-2 transmitted light microscope in
conjunction with QCapture Pro software. All samples, residues, and LM slides are curated in
the Centre for Palynology at the University of Sheffield, UK.

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Once a taxonomy had been established (at least one slide was logged for each sample
recording all taxa present) a minimum count was made for each sample. If sufficientpalynomorphs were present a minimum count of 200 was undertaken. When there wereinsufficient palynomorphs the entire palynomorph population was counted. To accommodate
for differences in quantitative approach, samples containing a yield greater than 200 wererarefied to 200. Damaged grains (>50%) identifiable as bisaccate or monosaccate pollen grains

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were logged as Unidentifiable to avoid misclassification of samples as barren. The count data
was recorded and manipulated using the packages StrataBugs 2.1 and R.

Depth bRT

SM11
Dove's
Nest

SM4
Gough

ZC LU

ZC LU

1250
1260

l{)

l!')

[:;j SS

N

1270

K4

Na4

1290

...

... A4

1300

w

Ca4

Upgang Fm.
Carnallitic
Marl Fm.

CM

SM7
Mortar
Hall

1340
1350

1400

w

~

Boulby Halite
Boulby Potash

Na3

ZC LU

Boulby Halite(")

Nw Billingham Anhydrite

Na3
(")

N

w

':

TD 1388m',[>- Litte Scar (QC)

Sneaton Halite
Sneaton Potash
Sneaton Halite
Sherburn Anhydrite

CM

1330

1390

N

Brockelschiefer
Littlebeck Anhydrite
Sleights Siltstone

Ca4

1320

1380

"<t"

w A4

N

N

131 0

1370

w

Na4

1280

1360

Triassic

A3
Brotherton Fm.

(")

N

w A3

141 0
1420

Ca3

1430

Grauer Salzton

SM14B
Wood smith
Mine

Ca3

ZC LU

GS

1440

GS

1450
1460

N

Rl
w FE

N

TD 1361m

TD 1360mw

1470

FordonEvaporite
Fm.

FE
N

N

w

1480
1490
1500
1510
N

N

1520

w

FE

Roker Dolomite/
Edlington Fm.

u
~

-

1530
1540

-

-

......

1560

Sp_r_otbrou_gh_
w Cadeby
Fm.

N

1570
1580

TD 1582m[:=] Anhydrite[:=] Halite[:=] Potash (K-Mg salts)[:=] Carbonates[:=] Continental/lacustrine sediments[:=] Terrestrial/red sandstone
-

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Hartlepool/Hayton
Anhydrite

1550

Conde nsed basinal Zechstein

I-

Marsden Bay (QC)
Levitt Haag Hole (QC)
Sandal House (QC)
Pot Riding (QC)
Ashfield Brick Pit (QC)
Kimberley (QC)

- Claxheugh Rock (QC),- Crime Rigg Quarry (QC)

Wetherby t - - Slaterford Farm (BH)
1--Marl Slate- Woolsthorpe Bridge (BH)
Rotliegend "yellow" sands

Brock = Brockelschiefer
A5 = Littlebeck Anhydrite
SS = Sleights Siltstone
Na4 = Sneaton (Halite)
K4 = Sneaton Potash
A4 = Sherburn Anhydrite
Ca4 = Upgang Fm.
CM = Carnallitic Marl Fm
Na3 = Boulby Halite

K3 = Boulby Potash
A3 = Billingham Anhydrite
Ca3 = Brotherton Fm.
GS = Grauer Salzton
FE = Fordon Evaporite Fm.
Ca2 = Edlington Fm.
Ca1b = Cadeby Fm. (Sprotbrough Member)
Ca1a = Cadeby Fm. (Wetherby Member)
MS = Marl Slate

Figure 3. Stratigraphy and correlation of the sequences analysed in this study. OC = Outcrop,
BH = borehole, ZC = Zechstein cycle, LU = Lithostratigraphic unit, and TD = Terminal Depth(m) of commercial boreholes. C = carbonates and E = evaporites.

269

5. Palynological results

270
271
272
273
274
275
276
277
278
279
280

Recovery was highly variable from the different lithologies sampled. Many of theclastic deposits yield abundant organic residue containing rich and diverse assemblages ofwell-preserved palynomorphs of low thermal maturity. Most of the evaporite deposits produced
very low yields of organic residue, although these often contained palynomorphs. However,palynomorphs from the evaporites were variable in preservation, varying both in quality ofpreservation (excellent to poor) and colour (translucent pale yellow to opaque black). All of
the palynological samples were dominated by pollen, with very rare spores, and only occasional
marine forms (acritarchs and foraminiferal test linings). A list of the taxa encountered isprovided in Table 1 and illustrations of these taxa are in Plates I-III. Figures 4-14 illustrate thepalynomorph occurrence and distribution for each locality. The raw data is provided inAppendix E. Table 2 summarises the palynomorph distribution.

281

Claxheugh RockSca le : 1:25

$1t(! (X)(le:Locat ion•

Sc aleCLR
54'54'28.47"N 1•26'24.02"'N

lnterval: 1.50

Lit0011tratigraphy

1.70m

_"_,

Palynology
SP

Sample Lit holosJy

l

I

C
D

c.

!w
282
283

J

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Tu o,, C~._;.o•■SPSj, o,M a,'o(l POll<)nl lFigure 4. Occurrence and distribution of palynomorphs from Claxheugh Rock.

Crime Rigg Quarry
Scale: 1 :25

Scale

f --

Site code:
LOCBtion:t -- - - -~CRQ
54°46'5.36~N 1D28'0 66"W
Lithostratigraphy- - - - -~lntcr,al: n.n - 4.0m

&ample Lithology- - - ---+- - - -- +- -~Palynology- - - - - - - -~S~P- - - - - - - - - - - - i BM-C Lithology■$h :!. leBoundary Typn

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0

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Q)

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284
285

Figure 5. Occurrence and distribution of palynomorphs from Crime Rigg Quarry.

Kimberley
Scale: 1:25

Sl1o co<1o· K1M
Location: 54°48'0.23"N 1' 13'8.76"WInterval: 3 93-4 23

Paly nology

Sa~ Llth.ology

SP_ _....·""""-•~..nd.,.,,.(200•00rrm. ......... "°' . 10 «1Yf'lb)

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ll I l

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286
287

i .

! i i. i t t I .!t I~

Figure 6. Occurrence and distribution of palynomorphs from Kimberley.
Salterford Farm
O\;::r:p~.: SF
CU•~
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288
289

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I

Figure 7. Occurrence and distribution of palynomorphs from Salterford Farm.

Woolsthorpe Bridge
Scale: 1 :25

Outcrop code : W B
Operator:
British Coal Board

Location· S4°4'492"N 1'5'5081"WInterval: 544m - 544m

Lithostratigra phy

Sample Litholo gy

Scale

Palynology
Base Uthok>gy

SP

I
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c.
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544.50

290
291
292

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!'
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~

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- - Confiden1
Samplin g-

cuttilg• Core..... Sidewal Core

Taxon Categories■

SP - Spores and pollen

"'

r-"

Figure 8. Occurrence and distribution of palynomorphs from Woolsthorpe Bridge.

Marsden Bay$(;;111• : 1:25

Siu. COO..

MB
54•51,•44_3,ij"N1" 23'37.51"\\'

293
294

Figure 9. Occurrence and distribution of palynomorphs from Marsden Bay.

SM4 GoughSeale: 1: 1000

Well Code: SM4GOUGH

Yor1c Potash Ltd

OperatOI':

Scale

LocatiOn: 54"2J'1 8.22"N 0"32'40.26"VV

1250m -1460m

Interval

Llthostratigraphy

Palynology

Lithology

8 - LMhology

SP■<,.,il<(l,;en'o-<)u;)<,l alN,I00,1()!(200 • &J)r'r~r, $(;! Q lo:;k • 10<»;nl$)

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Sherburn Anhydrite

1310,··,Uw><._r.,,-,..,..,.,

256

nm

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w:c

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Anhydrite-·.; ..':;:c·' ';: ;!;,•Brotherton
Formation

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EZ2

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I I --··••- ""• 1 'j1 .~

t/~()

)
'

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295
296
297

Figure 10. Occurrence and distribution of palynomorphs from SM4 Gough.

■~'••)'<ll~n~, ')i~■co1r-,..

!tll hal:Qii! _:;::,"9
•
■

N ~l

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299

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..... !!

r~

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en "I j301

Figure 11. Occurrence and distribution of palynomorphs from SM11 Dove’s Nest.

ISM14b Woodsmith Mine
Scale: 1:100

We i Code· SM 1ABWOOOSMITHMINE LOC11tioo· 54•2s· 11· N ooo•37'29"',','()p8r81.or: York Pol;,sh ltd
lnterv;,I
1442 m• 146 1m

e

"'

EZJ

Brolhllfton
For~ ~or,

Fordon Evaporite

format>on

302
303

Figure 12. Occurrence and distribution of palynomorphs from SM14b Woodsmith Mine.

SM7 Mortar Hall
Seale: 1:50

Woll Code: SM7MORTARHALL
Oper;:it or: York Potash Ltd

Seale

I
~

15.
~

"O

l lthostratlgr.iphy

.Q

1i

E

0

"-

Pa1yno1ogy

lithology

.ii<

1
~

LocatiOn: 54' 26'56.07"N 0°36'49.29"EInterval: 1382m 1388m

Bawlllhology

SP
QJ;'l~._-(!L0:1

I

ii

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f i~ ] ~I 'l}! ! !I ! i
1, '

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■

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Formation

G rauor Sal:i:ton

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Format,<>n

304
305
306

~)

••!-llf-S- c . ,Sampling

I

EZ3

EZ2

10e

BoundaryTn •e.

'Brotherton~- ~......J_~_l_,,..

1, m 1,s
M m~

••

l

"1i:

)

Figure 13. Occurrence and distribution of palynomorphs from SM7 Mortar Hall.

C\Jftir",j· w,-

S,,(1tw,)IC<:,rt

TH O<l C a ~ •■SP- Spc,-"51ndckm

Little Scar
Scale: 1:150

Site code:

Location:

LSSC
54°39'46 .S"N 1°11'12.0"W

Scale Lithostratigraphy Sample Lithology

Interval: 2.0 - 18.0m

Palynology
SP
·1

FOP

Text Keys•1 quantlsemi-quant abundan ..

·1

Base Litho logy
M mudstone (Dunham)■siltstone

Boundary Ty pes
Confident
~

E.
~

:J

iii

e
CJ

E
ro

i

roC -~~

16.
-

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16.0-.._,6.30

■

- ■

·

Taxo n Categories

<( ~

0.

<J)

■

core

- - Sidewall Core

en
>, !l

(/)
M
M
M

-

0

c.

a.

Cutting

.i

en
Q)

0
Q)

:i-

E.

C

Sampling

0.

<J)

g;

■

SP - Spores and pollen

"-

MEG
MEG
MEG

Brbckelschie fer- 14(P-T boundary)

12

10

Zechstein

307
308

Figure 14. Occurrence and distribution of palynomorphs from Little Scar.

309
310

5.1. General description of the spore/pollen assemblages

311
312
313
314
315

The assemblages are often dominated by the taeniate bisaccate Lueckisporites virkkiae(Plate I, 1-5) occurring at sufficient frequencies to dominate assemblages by ~50.0%. Variant
A (Plate I, 1, 2) is the most common variant, averaging 9.9-26.8% of assemblages, with Variant
B (Plate I, 3) and Variant C (Plate I, 4, 5) being noticeably less abundance, 1.7-5.1% and 1.59.4% respectively. L. virkkiae maintains dominance throughout the Zechstein sequence.

316
317
318
319
320
321
322
323
324

Taeniaesporites spp. (Plate I, 6-12), the most common species of which is T. labdacus(up to 14.0%, average 0.7-5.9%) (Plate I, 6, 7), is present up until the Cycle 4 carbonates but
no later. Average abundance ranges between 1.3-22.5%, with maximum abundance achievedin the Marl Slate. Protohaploxypinus spp. (Plate I, 13-16), the most common of which are P.
chaloneri (Plate I, 13, 14) and P. jacobii (Plate I, 15, 16), has a slightly greater temporal range
than Taeniaesporites, and is present into the Cycle 4 evaporites. Protohaploxypinus spp. ranges
in abundance from 0.6-2.2% of assemblages, but individual species can reach abundances of
3.4% (e.g. P. jacobii in the Cycle 3 evaporites). The presence and abundance of other taeniate
species is also as expected from Lopingian assemblages. The distinct asaccate species Vittatina

325
326
327
328
329

hiltonensis (Plate II, 1, 2) maintains a rare abundance throughout of 0.5-3.17%. Striatioabietites(Plate II, 3) and Striatopodocarpites (Plate II, 4, 5) occur at low abundances only, typically<0.5% of the assemblages, but up to 2.0% in the Cycle 3 evaporites. Platysaccus radialis (Plate
II, 6) is a rare bisaccate only occurring in the Cycle 1 carbonates (1.0%) and very rare in the
Cycle 4 carbonates (<0.5%).

330
331
332
333
334
335
336
337
338
339
340
341
342
343
344
345
346
347
348

Regarding non-taeniate bisaccate pollen, Klausipollenites schaubergeri (Plate II, 7, 8)
is the most abundant and is present from the Marl Slate through to the Cycle 4 evaporites. It issimilar in morphology to Vestigisporites minutus (Plate II, 9-11), but distinctly bisaccate rather
than monosaccate and larger in size. K. schaubergeri exhibits a large range in abundance but
on average represents between 5.8-20.0% of assemblages. A maximum abundance of 27.0% isachieved in the Cycle 2 carbonates, but abundances can be as low as 0.5%. Illinites spp. (Plate
II, 12-15) is the only other pollen grain taxon to occur in the Cycle 5 evaporites alongside
Lueckisporites virkkiae, however it is absent in the Cycle 4 evaporites and Cycle 5 carbonates.
It is easily identifiable with its distinct corpus and sacci shape meaning its presence can berecorded even in poorly preserved samples. Illinites spp. is represented by three species, with
I. delasaucei (Plate II, 12) being the most common (0.5-6.5%) and I. klausi (<0.5-3.5%) (Plate
II, 13) and I. tectus (0.5-5.2%) (Plate II, 14, 15) being present at lower abundances.
Falcisporites zapfei (Plate I, 12, 13) is the largest non-taeniate bisaccate recovered, occurring
at low abundances from the Marl Slate through to the Cycle 4 carbonates at <0.5-1.3%. The
distinctive bisaccate Alisporites nuthallensis (Plate II, 18) is present from the Marl Slatethrough to the Cycle 4 carbonates at low abundances of 0.5-2.4%. The triangular-outlined
pollen species Labiisporites granulatus (Plate II, 19, 20) was observed at average abundancesof 0.5-5.1%, but with abundances as high as 18.0% in the Cycle 3 evaporites. This species
exhibits one of the largest variations in abundance in the studied material.

349
350
351
352
353
354
355
356

Large monosaccate pollen species are, as expected, rare e.g., Nuskoisporites dulhuntyi(Plate III, 1), N. cf. rotatus (Plate III, 2), Perisaccus granulosus (Plate III, 3, 4) and
Potonieisporites novicus (Plate III, 5, 6). Nuskoisporites spp. and P. granulosus are present in
the Marl Slate through to the Cycle 4 carbonates, while Potonieisporites novicus has a similar
range yet is not present in the Marl Slate. These large monosaccate pollen typically only
represent 1.0-2.0% of assemblages or less. However, the small monosaccate Vestigisporites
minutus is one of the most common pollen species comprising on average 5.0% of assemblages.
V. minutus can comprise up to 17.5% of assemblages in Cycle 3 evaporites.

357
358
359
360
361
362
363
364
365
366

The possible cycad pollen, Cycadopites rarus, is very rare (0.5%). C. rarus is only
recorded in the Cycle 3 evaporites. Trilete spores (Plate II, 8, 9) maintain a low abundance
throughout Cycle 1-4 but reach their maximum presence of 13.6% in the assemblages of the
Cycle 4 evaporites. However, assemblages from Cycle 4 and Cycle 5 were of a lower yield
relative to earlier cycles. Trisaccate pollen grains are rare and may represent aberrant forms ofother pollen species (Foster and Afonin, 2005; Metcalfe et al., 2009). They occur at <0.5%
from Cycle 1 carbonates to Cycle 3 evaporites but are not found in assemblages beyond.
Tetrads are only found in Cycle 1 carbonates at maximum abundances of 0.5%. Cuticle
fragments of varying size (20-200µm) were common in assemblages, with larger fragments
recovered from the evaporite preparations than standard preparations.

367
368
369

5.2. Description of palynomorph distribution by locality

370
371

5.2.1. Cycle 1 transgressional sediments and carbonates (EZ1 Ca) (Marl Slate andcarbonates): Claxheugh Rock and Crime Rigg Quarry exposures

372
373
374
375
376
377
378

Palynomorph assemblages from Cycle 1 of the Durham Sub-basin were recovered fromexposures of the Marl Slate at Claxheugh Rock (Figure 4) and Crime Rigg Quarry (Figure 5).
Palynological preparations contain vast amounts of amorphous organic matter (AOM).
Palynomorphs are present and are well-preserved and hyaline in appearance. The hyaline
nature hampered identification of palynomorphs, but genus-level identifications, and somespecies-level identifications of distinct taxa, were possible after samples were stained with
Bismarck Brown.

379
380
381
382
383
384
385
386
387
388

The assemblages recovered from the exposure at Claxheugh Rock (Figure 4) werecomposed of taeniate bisaccate species: Lueckisporites virkkiae (31.5%), Protohaploxypinus
sp. (1.0%), Taeniaesporites sp. (17.0%) and Vittatina hiltonensis (1.0%). Non-taeniate
bisaccate species include Alisporites nuthallensis (<0.5%), Illinites sp. (2.0%), Klausipollenites
schaubergeri (8.5%), rare Labiisporites granulatus (<1.0%) and Falcisporites zapfei (<0.5%).
Rare multisaccate Crustaesporites globosus (<0.5%) (Plate III, 7) was present. Monosaccate
pollen was represented by Vestigisporites minutus (3.5%) and rare Perisaccus granulosus(<0.5%) and Nuskoisporites dulhuntyi (0.5%). One sample (CLR) did not contain
Unidentifiable palynomorphs but contains rare unidentifiable spores (<0.5%), while the othersample (CLR2) contains 93.0% Unidentifiable palynomorphs.

389
390
391
392
393
394

The assemblage from Crime Rigg Quarry (Figure 5) was composed of the taeniate
bisaccate species Lueckisporites virkkiae (43.5%), Protohaploxypinus sp. (2.5%), rare
Striatopodocarpites sp. (<0.5%%), Taeniaesporites sp. (28%), and Vittatina hiltonensis(2.0%), and the non-taniate bisaccate species Illinites sp. (7.0%), Klausipollenites schaubergeri(8.0%), Labiisporites granulatus (1.5%), Falcisporites zapfei (<0.5%), as well as themonosaccate pollen Perisaccus granulosus (<0.5%) and Vestigisporites minutus (7.5%).

395
396
397
398

5.2.2. Cycle 1 transgressional sediments and carbonates (EZ1 Ca) (Cadeby Fm.:
Wetherby/Sprotbrough Mb.): Kimberley railway cutting exposure and the Salterford Farm andWoolsthorpe Bridge boreholes

399
400
401
402
403
404
405
406
407
408
409
410

Palynomorph assemblages from Cycle 1 of the Yorkshire Sub-basin were recoveredfrom the Cadeby Fm. exposure in Kimberley railway cutting (Figure 6), Salterford Farm
borehole (Figure 7), and Woolsthorpe Bridge borehole (Figure 8). The palynomorphs were
well preserved and translucent. Both assemblages from Kimberley railway cutting, K5 and K6,
contain Lueckisporites virkkiae (K5 39.0%, K6 50.0%), Taeniaesporites sp. (K5 2.5%, K6
1.0%), T. labdacus (K5 10.5%, K6 6.5%), T. novimundi (K5 2.5%, K6 2%), T. noviaulensis(K5 2.5%, K6 5.5%), T. angulistriatus (K5 2.0%, K6 2.0%), T. albertae (K6 7.5%),
Protohaploxypinus jacobii (K5 2.0%), Striatopodocarpites antiquus (K5 1.0%, K6 0.5%),
Vittatina hiltonensis (K5 2.5%), Illinites sp. (K5 5.5%), I. delasaucei (K5 9.5%, K6 8.5%), I.
klausi (K6 3.5%), I. tectus (K5 1.0%, K6 0.5%), rare Nuskoisporites dulhuntyi (K5 <0.5%) and
Potonieisporites novicus (K5 <0.5%), rare Perisaccus granulosus (K6 <0.5%), unidentifiable
spores (K5 2%, K6 0.5%), and rare tetrads (K6 0.5%).

411
412
413
414
415

Assemblages from the Salterford Farm borehole samples were composed of the taeniate
bisaccate species Lueckisporites virkkiae with Variant A (21.3%), Variant B (1.7%) and
Variant C (2.5%), Protohaploxypinus sp. (0.5%) P. chaloneri (<0.5%), P. jacobii (0.5%),
Striatoabieites richteri (0.5%), Striatopodocarpites sp. (0.5%), Taeniaesporites sp. (0.5%), T.
albertae (0.8%), T. angulistriatus (0.8%), T. bilobus (<0.5%), T. labdacus (2.8%), T.

416
417
418
419
420
421
422
423
424
425

novimundi (1.3%), T. noviaulensis (0.8%). The non-taeniate bisaccate pollen Alisporites
nuthallensis (2.5%), Falcisporites zapfei (0.9%), Illinites delasaucei (5.1%), I. tectus (0.5%),
Klausipollenites schaubergeri (14.3%), Labiisporites granulatus (3.5%), Platysaccus radialis(1.0%) and Potonieisporites novicus (0.5%) were recovered. The multisaccate Crustaesporites
globosus was recorded (0.5%). Monosaccate pollen were represented by Nuskoisporites
dulhuntyi (0.5%), Perisaccus granulosus (0.6%) and Vestigisporites minutus (6.7%).
Reduviasporonites sp. was found throughout (2.7%), trisaccates were rare (0.4%) and spores
were found in all samples (1.6%). A solitary tetrad in YFP6368 was recovered and a single
foraminiferal test lining was recovered from SF465. The Unidentifiable palynomorph
component averages at 33.0%.

426
427
428
429
430
431

The sample of Marl Slate obtained from Woolsthorpe Bridge borehole (Figure 8) was
not species rich and was dominated by Unidentifiable palynomorphs (56.1%). Identifiabletaeniate bisaccate pollen include Lueckisporites virkkiae Variant A (15.8%), Variant B (1.8%)
and Variant C (3.5%), Taeniaesporites labdacus (1.8%), T. albertae (3.5%), and non-taeniate
bisaccate pollen include Klausipollenites schaubergeri (12.3%) and Labiisporites granulatus(1.8%). The only monosaccate taxon present was Vestigisporites minutus (3.5%).

432
433
434
435

The remaining Cycle 1 carbonates, sampled at Ashfield Brick Pit, Sandal House, and
Pot Riding, were essentially barren of palynomorphs, containing only a few palynomorphs ofsuch poor preservation they lacked characters that enabled them to be identified even as
Unidentifiable pollen grains. Therefore, these sites were designated as barren.

436
437

5.2.3. Cycle 1 evaporites (EZ1 A) (Hartlepool Anhydrite/Hayton Anhydrite)

438
439
440

Cycle 1 evaporites were not covered in the sampling range of this study as they are notrepresented by any of the borehole material and they are not exposed at outcrop as a result of
dissolution.

441
442
443

5.2.4. Cycle 2 carbonates (EZ2 Ca) (Roker Dolomite/Edlington Fm. and Kirkham Abbey Fm.):
Marsden Bay

444
445
446
447
448
449
450
451
452
453
454
455
456
457
458
459
460

A sample of red mudstone from the Edlington Fm. collected from outcrop at Levitt
Hagg Hole was barren of palynomorphs. Samples of Roker Dolomite Fm. from Marsden Bay(MB1-6) (Figure 9) contained taeniate and non-taeniate bisaccate pollen, as well as raremonosaccate pollen. The assemblage of MB2 was the best preserved. The taeniate bisaccate
taxa were represented by Lueckisporites virkkiae (53.5%), and Taeniaesporites sp. (1.8%) is
also abundant, with T. angulistriatus (<0.5%), T. labdacus (0.5%) and T. novimundi (<0.5%).
Protohaploxypinus sp. (3.1%) was present, with P. chaloneri (<0.5%) and P. jacobii (<0.5%)
present. Striatopodocarpites sp. (<0.5%) was rare and only found in one sample (MB2).
Vittatina hiltonensis (0.9%) was present at rare abundance in MB1-3. Smooth bisaccate pollen
was represented by abundant Illinites sp. (6.1%) with rare I. delasaucei (<0.5%) only recoveredfrom MB2. Klausipollenites schaubergeri was abundant (23.3%), while Labiisporites
granulatus (1.0%), Alisporites nuthallensis (0.9%) was uncommon, and Falcisporites zapfei(<0.5%) (MB4-6) and Potonieisporites novicus were rare (<0.5%). The monosaccate pollen
Nuskoisporites dulhuntyi, and Perisaccus granulosus were rare (<0.5%), and Vestigisporites
minutus (MB1-2) was found in MB1-2. The Unidentifiable component was only present in
MB1 and only comprises 16% of the assemblage. Unidentifiable trisaccate pollen grains wererecovered from MB1 (<0.5%).

461
462
463
464
465
466
467
468
469
470
471

The remaining samples from Marsden Bay (MB4-6) contained similar assemblages, but
typically of lower yield. MB4 contains an exceptionally well-preserved assemblage but was of
low yield (n=102), with Lueckisporites virkkiae composing 20.0% of the assemblage. There
were no Unidentifiable palynomorphs, but L. virkkiae Variant A was abundant (14.0%) while
Variant B was rare (<0.5%). Other taeniate genera Taeniaesporites sp. were present (1.8%) as
well as Protohaploxypinus spp. which was only found in MB4 (0.5%), with rare P. chaloneri(<0.5%) and P. microcorpus (<0.5%) recovered from MB5. The non-taeniate bisaccate pollen
Illinites sp. (10.7%), Falcisporites zapfei (0.5%), and Klausipollenites schaubergeri (16.7%)
were abundant in MB4-6. The only monosaccate taxon recovered from MB4-6 was the small
species Vestigisporites minutus that was only recovered from MB5 (2.5%). Unidentifiable
pollen grains comprised on average 54.5% of the assemblages.

472
473
474

5.2.5. Cycle 2 evaporites (EZ2 A, EZ2 K, EZ2 Na) (Fordon Evaporites Fm): SM4, SM11 andSM14b boreholes

475
476
477
478
479
480

The borehole material captures both the lower (SM11) and upper (SM4, SM14b) partsof the Fordon Evaporite Fm.. In borehole SM11 (Figure 10), the lower part of the Fordon
Evaporite Fm. contained assemblages with a low yield. Palynomorphs were mostly of the
Unidentifiable type, corroded, dark and fragmented. However, some very well-preserved
specimens of Illinites sp., Klausipollenites schaubergeri, Reduviasporonites sp.,
Lueckisporites virkkiae Variant A and Variant B, and Vestigisporites minutus were recovered.

481
482
483
484
485
486

The upper part of the Fordon Evaporite Fm. in SM11 (Figure 10) contained a morevaried assemblage with samples containing greater yields. Very high proportions of
Unidentifiable pollen grains persisted, as well as some very well-preserved palynomorphs
originating in the evaporites. Assemblages were dominated by Lueckisporites virkkiae Variant
A, Klausipollenites schaubergeri, Vestigisporites minutus and assorted Illinites sp.,
Taeniaesporites sp. and Protohaploxypinus sp.

487
488
489
490
491
492
493
494
495
496
497
498
499

On average assemblages from the Fordon Evaporite Fm. from SM11 (Figure 10)
contained taeniate bisaccate species Lueckisporites virkkiae Variant A (13.3%), as well as
Variant B (4.1%) and Variant C (3.8%), Taeniaesporites sp. (5.2%), rare T. angulistriatus(0.5%), T. albertae (1.3%), T. noviaulensis (1.0%), and T. labdacus (1.31%%), rare
Protohaploxypinus sp. (1.3%) including P. chaloneri (0.8%), P. jacobii (0.5%) and P.
microcorpus (0.5%). Striatopodocarpites antiquus (0.5%) and Vittatina hiltonensis (0.5%)
were very rare. The non-taeniate bisaccate pollen were represented by Klausipollenites
schaubergeri (6.4%), Illinites sp. (5.1%) including I. delasaucei (1.8%), and rare Labiisporites
granulatus (1.8%), Falcisporites zapfei (0.5%) and Potonieisporites novicus (0.5%).
Monosaccate pollen were rare and were represented by Nuskoisporites dulhuntyi (0.5%), yet
Vestigisporites minutus was abundant (6.7%). Reduviasporonites sp. was present (9.6%), yet
samples were dominated by Unidentifiable palynomorphs (74.52%). Some trisaccate pollen
grains were recovered (6.67%), as well as rare spores (0.5%) and very rare acritarchs (<0.5%).

500
501
502
503

The Unidentifiable component appears to increase in prominence throughout the upper
part of the Fordon Evaporite Fm. Species richness and abundance of palynomorphs also
appears to increase throughout the Fordon Evaporite Fm. as the proportion of Unidentifiable
palynomorphs progressively dominates samples.

504
505
506

Borehole SM4 (Figure 11) captures the top of the Fordon Evaporite Fm., where samples
either had a high yield or a very low yield (<100 pollen grains) yet were still dominated by
Unidentifiable palynomorphs (95.1%), which indicates pollen transport from a nearby source

507
508
509
510
511

vegetation. In addition to the Unidentifiable palynomorphs, rare Lueckisporites virkkiae (1.8%) including Variant A (0.5%) and L. virkkiae Variant C (1.5%), Taeniaesporites sp. (4.3%),
Protohaploxypinus chaloneri (1.5%), Klausipollenites schaubergeri (0.5%) and Vittatina
hiltonensis (<0.5%), and unidentifiable spores (1.0%) were recovered. Only in the lower partsof the Fordon Evaporite Fm. were the three “Variants” of L. virkkiae distinguishable.

512
513
514
515
516
517

In borehole SM14b (Figure 12), the Fordon Evaporite Fm. palynomorph assemblages
were generally well-preserved and of a high yield. Although containing a high proportion of
Unidentifiable palynomorphs (80.0%), Lueckisporites virkkiae Variant A (11.0%) and Variant
C (5%) are present, as well as Klausipollenites schaubergeri (5.9%), Potonieisporites novicus(7.0%) and Illinites delasaucei (7.0%), and Reduviasporonites sp. (3.0%). L. virkkiae Variant
C was the latest variant to appear in the core relative to the other two.

518
519
520

5.2.6. Cycle 2 – Cycle 3 transitional (EZ3 Ca) (grey salt clay/illitic shale/Grauer Salzton):
SM4 borehole

521
522
523
524
525
526
527
528
529
530

In SM4 (Figure 11) the Grauer Salzton/Brotherton Fm. boundary assemblage wasdominated by Unidentifiable palynomorphs (87.7%), but unlike the other boreholes containing
palynomorphs identifiable to species level the assemblage was found to contain Lueckisporites
virkkiae Variant A (3.8%), Variant B (0.9%), and Variant C (2.8%), Illinites sp. (0.9%),
Labiisporites granulatus (1.9%), Taeniaesporites sp. and T. labdacus (0.9%). In borehole SM7(Figure 13) a single sample captures the palynology of the Grey Salt Clay. It contains anabundant assemblage composed only of Unidentifiable palynomorphs (100.0%). In borehole
SM11 (Figure 10) this transitional stratum was either barren, or was composed of a very
abundant sample of only 100.0% Unidentifiable palynomorphs. In SM14b (Figure 12)
assemblages of the Grauer Salzton are barren.

531
532

5.2.7. Cycle 3 carbonates (EZ3 Ca) (Brotherton Fm.): SM7, SM11 boreholes

533
534
535
536
537
538
539
540

In borehole SM7 (Figure 13) a single assemblage from the Brotherton Fm. wasrecovered. Though of low yield (<200) the assemblage contains the taeniate bisaccate species
Lueckisporites virkkiae (10.0%), including “Variants A” (2.5%), Variant B (1.0%) and veryrare Variant C (0.5%). Taeniaesporites sp. (1.0%) was rare with T. noviaulensis (<0.5%) being
very rare. Protohaploxypinus sp. was rare (<0.5%), Illinites sp. (1.5%) is present.
Klausipollenites schaubergeri (8.0%) was present, Vestigisporites minutus was rare (0.5%) andwas the only monosaccate species recovered. Well over half of the assemblage was composedof Unidentifiable palynomorphs (74.0%).

541
542
543
544
545
546
547

Many of the Brotherton Fm. samples from borehole SM11 (Figure 10) were barren(n=15). Recovered assemblages contain abundant palynomorphs, but were dominated by
Unidentifiable palynomorphs (90.2%), presumably an effect of adverse preservational
conditions during deposition. There appears to be an increase in the Unidentifiable component
from the end of Cycle 2 evaporites through to the Brotherton Fm. However, not all samples
were dominated by Unidentifiable palynomorphs. Some assemblages were very wellpreserved.

548
549
550
551

Pollen taxa that occured in low abundances include the taeniate bisaccate species
Lueckisporites virkkiae (4.8%) including abundant L. virkkiae Variant A (10.4%), with L.
virkkiae Variant B (3.5%) and Variant C (1.3%), Protohaploxypinus sp. (3.3%),
Striatopodocarpites antiquus (0.8%), Taeniaesporites spp. (2.0%) including T. angulistriatus

552
553
554
555
556
557
558

(0.5%), T. noviaulensis (0.5%), and T. labdacus (0.5%). Non-taeniate bisaccate pollen
recovered were Alisporites nuthallensis (<0.5%), Falcisporites zapfei (<0.5%), Illinites sp.
(3.3%) I. klausi (<0.5%) and I. delasaucei (0.5%), Klausipollenites schaubergeri (5.9%),
Labiisporites granulatus (1.0%) and Potonieisporites novicus (<0.5%). The monosaccate
pollen Vestigisporites minutus (7.8%), Nuskoisporites cf. rotatus (<0.5%), and Nuskoisporites
dulhuntyi (<0.5%) were present. Trisaccate pollen grains and spores, and Reduviasporonites
sp. were all rare (<0.5%).

559
560
561
562

The sample of Brotherton Fm. from SM4 (Figure 11) was barren. However, the sampleof Brotherton Fm. from SM14b (Figure 12) contained an abundant yield of pollen, dominatedby Unidentifiable palynomorphs (99.5%), but also contained very rare occurrences of
Lueckisporites virkkiae (1.7%), and Taeniaesporites sp. (0.6%)

563
564
565

5.2.8. Cycle 3 evaporites (Billingham Anhydrite (EZ3 A), Boulby Potash (EZ3 K) and Boulby
Halite (EZ3 Na)): SM11 borehole

566
567

Assemblages in borehole SM11 (Figure 10) became better-preserved throughout theevaporites, especially in the Billingham Anhydrite and Boulby Halite.

568
569

5.2.9 Cycle 3 anhydrite (Billingham Anhydrite): SM11 borehole

570
571

There was no recovery from the Billingham Anhydrite or lower part of the Boulby
Halite in SM4 (Figure 11).

572
573
574
575
576
577
578
579
580
581
582
583
584

In SM11 (Figure 10) the Billingham Anhydrite assemblages yielded Lueckisporites
virkkiae (7.5%), including Variant A (16.9%), Variant B (2.8%) and Variant C (3.5%),
Protohaploxypinus sp. (1.3%), P. chaloneri (1.3%), P. jacobii (1.4%), P. microcorpus (0.9%)
and P. cf. samoilovichii (0.5%), Taeniaesporites sp. (2.8%), T. angulistriatus (1.2%), T.
labdacus (2.6%) and T. novimundi (1.5%) and T. noviaulensis (1.5%), Striatopodocarpites sp.
(0.6%), Striatoabieites antiquus (1.6%) and rare Vittatina hiltonensis (<0.5%). Non-taeniate
bisaccate pollen were represented by Alisporites nuthallensis (1.8%), Falcisporites zapfei(0.9%), Illinites sp. (5.0%), I. delasaucei (4.2%), Klausipollenites schaubergeri (16.3%),
Labiisporites granulatus (1.5%), Potonieisporites novicus (0.8%). The multisaccate
Crustaesporites globosus was present but rare (0.5%). The monosaccate pollen wererepresented by Nuskoisporites dulhuntyi (0.8%), Perisaccus granulosus (0.9%) andVestigisporites minutus (6.3%). Reduviasporonites sp. (1.7%), spores (1.8%), and acritarchs(0.5%), were rare.

585
586

5.2.10. Cycle 3 potassium-magnesium salts (Boulby Potash): SM11 borehole

587
588
589
590
591
592
593
594

In SM11 (Figure 10) the Boulby Potash yielded an assemblage composed of the taeniate
bisaccate species Lueckisporites virkkiae (49.5%), Protohaploxypinus sp. (0.5%), P. jacobii(2.5%), P. microcorpus (1.5%), Taeniaesporites albertae (2%), T. labdacus (2.5%) and T.
noviaulensis (2.0%), the non-taeniate bisaccate species Falcisporites zapfei (1.0%), Illinites
tectus (4.0%), Klausipollenites schaubergeri (4.5%), Labiisporites granulatus (18.0%),
Potonieisporites novicus (0.5%), and the monosaccate species Vestigisporites minutus (5.0%)
and Perisaccus granulosus (1.0%). Reduviasporonites sp. was present (5.0%), as well as veryrare spores (<0.5%).

595
596

5.2.11. Cycle 3 halite (Boulby Halite): SM4, SM11 boreholes

597
598
599
600
601
602
603
604
605
606

In borehole SM11 (Figure 10) the Boulby Halite contained the taeniate bisaccate
species Lueckisporites virkkiae (13.0%) including Variant A (21.7%), Variant B (1.4%) and
Variant C (9.3%), Protohaploxypinus sp. (0.5%), including the species P. chaloneri (2.3%) andP. jacobii (2.5%), Taeniaesporites sp. (3.76%), T. angulistriatus (0.5%), T. labdacus (2.5%),
the non-taeniate bisaccate species Falcisporites zapfei (1.0%), Illinites sp. (<0.5%),
Klausipollenites schaubergeri (8.9%), Labiisporites granulatus (9.3%), Potonieisporites
novicus (0.5%). The monosaccate pollen were represented by Vestigisporites minutus (6.2%)
and Perisaccus granulosus (1.0%). Reduviasporonites sp. was present (2.6%) as well as veryrare trisaccate species (<0.5%). The Unidentifiable component averaged at 58.7% of theassemblage.

607
608
609
610
611
612
613

In borehole SM4 (Figure 11) assemblages of Boulby Halite were either barren or
contained assemblages of low yield, dominated on average 90.8% by Unidentifiable
palynomorphs. Klausipollenites schaubergeri (8.7%), Lueckisporites virkkiae (11.0%),
Vestigisporites minutus (2.9%), Taeniaesporites sp. (1.45%) including T. novimundi (2.9%)
were recovered. It is in the Boulby Halite that Vestigisporites minutus makes its first
appearance in borehole SM4, although earlier occurrences cannot be excluded on the basis ofrecovery of V. minutus in the Fordon Evaporite Fm. of borehole SM11.

614
615

5.2.12. Cycle 3 – Cycle 4 transition Carnallitic Marl Fm. (Rotten Marl): SM4, SM11 boreholes

616
617
618

Exceptionally well-preserved assemblages were extracted from the Carnallitic Marl
Fm. in borehole SM11 (Figure 10) using standard palynological acid maceration techniques,representing the first time palynomorphs have been recorded from the formation.

619
620
621
622
623

In SM4 (Figure 11) an assemblage from the base of the Carnallitic Marl Fm. was notdiverse or of high yield, but it was sufficiently well-preserved, relative to the underlying
Boulby Halite, to recover Lueckisporites virkkiae (11.6%), Klausipollenites schaubergeri(8.7%), Vestigisporites minutus (9.2%), Taeniaesporites sp. (1.5%) including Taeniaesporites
novimundi (2.9%), and Unidentifiable palynomorphs (72.5%)

624
625
626
627
628
629
630
631
632
633
634
635
636
637
638
639

Assemblages from the Carnallitic Marl Fm. of borehole SM11 (Figure 10) were ofvarying quality. Assemblages contained moderate amounts of Unidentifiable palynomorphs(15.3%) as well as distinctly well-preserved pollen grains. Taeniate bisaccate pollen wererepresented by the species Lueckisporites virkkiae (52.0%) including L. virkkiae Variant A(20.0%), L. virkkiae Variant B (4.4%), L. virkkiae Variant C (3.6%), Protohaploxypinus sp.
(1.5%), P. chaloneri (1.7%), P. jacobii (0.8%), Striatopodocarpites fusus (<0.5%),
Taeniaesporites sp. (3.0%), T. albertae (1.3%), T. angulistriatus (2.7%), T. bilobus (0.5%), T.
labdacus (5.9%), T. noviaulensis (2.2%), T. novimundi (0.6%), and Vittatina hiltonensis(0.6%). Non-taeniate bisaccate pollen were represented by Alisporites nuthallensis (0.8%),
Falcisporites zapfei (1.2%), Illinites sp. (3.0%), I. delasaucei (2.5%), I. klausi (0.5%), I. tectus(2.1%), Klausipollenites schaubergeri (11.4%), Labiisporites granulatus (0.8%),
Potonieisporites novicus (0.5%). The multisaccate Crustaesporites globosus was present(1.0%). Monosaccate pollen were represented by Nuskoisporites dulhuntyi (<0.5%),
Perisaccus granulosus (0.5%), and Vestigisporites minutus (5.1%). Reduviasporonites sp.
(2.5%) and rare acritarchs (0.5%) were also present. The Carnallitic Marl Fm. in SM11 is the
only recorded instance of Taeniaesporites bilobus in borehole SM11.

640
641

5.2.13. Cycle 4 carbonates (EZ4 Ca) Upgang Fm.: SM4, SM11 boreholes

642
643
644
645
646
647
648
649
650
651

In SM11 (Figure 10) the Upgang Fm. yielded assemblages composed of the taeniate
species Lueckisporites virkkiae (52.0%), Protohaploxypinus chaloneri (0.5%), P. jacobii(<0.5%), Taeniaesporites angulistriatus (1.0%), T. labdacus (6.0%), T. noviaulensis (1.5%),
T. novimundi (1.0%), Striatoabieites richteri (<0.5%), Striatopodocarpites antiquus (<0.5%),
Vittatina hiltonensis (1.0%), and the non-taeniate bisaccate species Falcisporites zapfei (1%),
Illinites delasaucei (2.0%), Klausipollenites schaubergeri (16.5%), Labiisporites granulatus(1.5%), Potonieisporites novicus (<0.5%). The monosaccate pollen were represented by
Perisaccus granulosus (0.5%) and Vestigisporites minutus (9.0%). Reduviasporonites sp.
(1.0%) was present but rare, and the Unidentifiable component was comparatively reduced(4.0%). The Upgang Fm. was not sampled in SM4 (Figure 11).

652
653
654

5.2.14. Cycle 4 evaporites (Sherburn Anhydrite (EZ4 A), Sneaton Halite (EZ4 Na) and Sneaton
Potash (EZ4 K)): SM4, SM11 boreholes

655
656
657
658
659
660

Analysis of rock salt samples yielded palynological recovery from the Cycle 4
evaporites. Palynomorphs have not previously been recorded from either the Cycle 4 or Cycle
5 evaporites as they are not lithologies typically targeted for palynological analysis and areoften missing from borehole cores as the underlying Carboniferous and earlier Permian strata
are more commercially important. Therefore, the results presented here from Cycle 4 and above
provide a unique insight into the vegetation during latest Zechstein times.

661
662
663
664

In SM11 (Figure 10) two samples from the lower Sneaton Halite, were of low yield andcomposed mostly of Unidentifiable pollen grains, however Lueckisporites virkkiae Variant C,
Klausipollenites schaubergeri, and Reduviasporonites sp. were recovered as well. In SM11
one sample from the upper Sneaton Halite contained only rare Unidentifiable pollen (n=3).

665
666
667
668
669

In SM11 the Sherburn Anhydrite assemblage was dominated by Unidentifiable pollen
grains (71.4%), and contained the taeniate bisaccate pollen Lueckisporites virkkiae (3.2%), L.
virkkiae Variant C (9.4%), the non-taeniate bisaccate pollen Klausipollenites schaubergeri(1.6%), Labiisporites granulatus (9.1%), Potonieisporites novicus (9.1%), and considerable
abundance of Reduviasporonites sp. (22.8%).

670
671
672
673
674

In borehole SM4 (Figure 11), the Sneaton Halite and Sneaton Potash assemblages did
not have a high yield (<200 count) and are dominated by Unidentifiable palynomorphs (88.9100.0%), though they contained Klausipollenites schaubergeri (7.4%) and Lueckisporites
virkkiae (3.7%). The upper Sneaton Halite of borehole SM11 contained a single sample of very
low yield composed only of Unidentifiable palynomorphs.

675
676
677

The Sneaton Potash in SM4 was dominated by Unidentifiable pollen grains (91.8%),
with the assemblage also containing Lueckisporites virkkiae (8.2%) and Klausipollenites
schaubergeri (8.5%).

678
679

5.2.15. Cycle 5 carbonates (EZ5 Ca) Sleights Siltstone: SM4 borehole

680
681
682

The Sleights Siltstone from borehole SM4 (Figure 11) contained an assemblage
composed only of Unidentifiable palynomorphs at a very low yield (<200 count) (sample SM41258.63 m).

683
684

5.2.16. Cycle 5 evaporites (Littlebeck Anhydrite and Bröckelschiefer): SM11 borehole

685
686
687
688

There was no recovery from the Littlebeck Anhydrite in any of the borehole material.
However, the Bröckelschiefer in SM11 (Figure 10) yielded an assemblage composed of
Lueckisporites virkkiae Variant C (2.0%), Illinites sp. (5.0%) and dominated by Unidentifiable
pollen grains (93.0%)

689
690

5.2.17. Latest Zechstein: Little Scar Beach outcrop

691
692
693
694
695
696
697
698
699
700

Samples from Little Scar (Figure 14) were mostly barren yet those that did contain ayield had a very low abundance of palynomorphs (<50). However, Taeniaesporites sp. wasfound in LSSC 1 (2.5%), and LSSC 1-6 did contain very low abundances of Unidentifiable
palynomorphs (80.0-100.0%). LSSC 1 had the highest yield of palynomorphs at 38
palynomorphs per slide. What is most interesting about these samples is that LSSC 2 and LSSC
3 contained benthic foraminiferal test linings (Plate III, 10), which are indicative of relatively
normal marine conditions. Foraminiferal test linings were incredibly rare throughout thematerial used in this study, only otherwise found in material from Salterford Farm boreholebelonging to the Marl Slate (EZ1) (Plate III, 11). Belonging to Cycle 4-5, the strata at Little
Scar beach form a useful comparison the strata of a younger age from Cycle 1.

701
702

5.3. Summary of the distribution of taxa within the stratigraphical sequence

703
704

5.3.1. Cycle 1 Marl Slate: the principal Zechstein transgression

705
706
707
708
709
710
711
712
713
714
715

The Marl Slate assemblages are generally very well preserved as they were recoveredfrom bituminous anoxic shale. In terms of average percentage abundance, they are dominatedby Lueckisporites virkkiae (53.5%) and Taeniaesporites spp. (22.5%), but also contain
Klausipollenites schaubergeri (8.3%), Illinites sp. (5.5%), Vestigisporites minutus (5.5%),
Vittatina hiltonensis (2.5%), and less than 2.0% each of Alisporites nuthallensis,
Crustaesporites
globosus,
Nuskoisporites
dulhuntyi,
Perisaccus
granulosus,
Protohaploxypinus spp., Striatopodocarpites sp. and spores. In samples where Unidentifiable
pollen grains are present, they represent 93.0% of the assemblage indicating poor preservation.
Many pollen grains are of a hyaline nature making species identification difficult. Recovery ofbenthic coiled foraminiferal test linings from Salterford Farm borehole sample SF465
corroborates marine conditions during deposition.

716
717

5.3.2. Cycle 1 Carbonates

718
719
720
721
722
723
724
725

The Cycle 1 carbonates assemblages are generally more speciose than those from theMarl Slate and the palynomorphs are less hyaline in nature making species differentiation andidentification easier. Assemblages are still dominated by Lueckisporites virkkiae (44.5%), buthere the three variants of Clarke (1965) are distinguishable. Klausipollenites schaubergeri ismore abundant than in the Marl Slate (13.7%) as well as Illinites sp. (5.5%) but with the three
species I. delasaucei, I. klausi and I. tectus distinguishable, of which I. delasaucei reachedabundances of 9.5%. Vestigisporites minutus is of similar abundance (4.8%) while Vittatina
hiltonensis increased in abundance (3.2%). A variety of Taeniaesporites spp. are present

726
727
728
729
730
731
732
733
734
735
736

including T. albertae (5.5%), T. angulistriatus (1.7%), T. labdacus (4.5%), T. noviaulensis(3.0%) and T. novimundi (1.8%). Alisporites nuthallensis (2.4%) and Labiisporites granulatus(2.6%) increase in abundance. Protohaploxypinus sp. (0.6%), P. jacobii (1.3%), andFalcisporites zapfei (1.0%) are less abundant than in the Marl Slate, becoming rare, while
Nuskoisporites dulhuntyi (0.5%), Perisaccus granulosus (0.5%), and Striatopodocarpites sp.
(0.5%) maintain a rare abundance. Striatoabieites richteri (0.5%) and Reduviasporonites sp.
(3.4%) appear, along with very rare tetrads (<0.5%). Crustaesporites globosus, Cycadopites
rarus and acritarchs are not present. In the Cycle 1 carbonates better preservation enables theidentification of more species, giving the impression of increased species richness. According
to previous literature Zechstein Cycles 1 and 2 yielded the richest palynomorph assemblages,
which this data supports, but a preservational bias should not be ignored.

737
738

5.3.3. Cycle 2 Carbonates

739
740
741
742
743
744
745
746
747
748
749
750
751
752
753
754

The Cycle 2 carbonate assemblages are also dominated by Lueckisporites virkkiae(53.5%), but only Variant A is distinguishable. Alisporites nuthallensis becomes rare (1.0%),
Illinites sp. increases in abundance (8.4%), I. delasaucei decreases dramatically (<0.5%) andI. klausi and I. tectus disappeared. Klausipollenites schaubergeri (20.0%) and Taeniaesporites
spp. (4.6%) increases in abundance yet fewer species are present with T. noviaulensis absent,
and overall the abundance of individual species decreases, suggesting Taeniaesporites is notwell enough preserved to distinguish species in these assemblages. Protohaploxypinus spp.
continues to be rare (2.2%) however more species are present including P. chaloneri (0.5%),
P. jacobii (<0.5%), and P. microcorpus (1.0%). Nuskoisporites dulhuntyi (1.0%) andPerisaccus granulosus (<0.5%) maintains a rare abundance. Vestigisporites minutus decreases
in abundance by over a half (1.9%). Falcisporites zapfei (1.0%), Vittatina hiltonensis (0.5%),
spores (<0.5%), Striatopodocarpites sp. (<0.5%) and Potonieisporites novicus (<0.5%) are
rare. Striatoabieites richteri, Platysaccus radialis, and Reduviasporonites sp. disappears andCrustaesporites globosus, Cycadopites rarus, acritarchs, and tetrads are still absent.
Unidentifiable pollen grains comprises on average 44.9% of assemblages, slightly less (~4.5%)
than in the Cycle 1 carbonates.

755
756

5.3.4. Cycle 2 Evaporites

757
758
759
760
761
762
763
764
765
766
767
768
769
770
771
772

The Cycle 2 evaporite assemblages contains Lueckisporites virkkiae (5.5%) at much
lower abundances but with all three variants distinguishable, due to a relative increase in theabundance of Unidentifiable pollen grains (77.4%), a result of many poorly preserved
assemblages. Consequently, the abundance of many species appears to decline. Illinites sp.
experiences a slight decline in abundance (5.6%) with I. delasaucei increasing in abundance(2.5%), and I. klausi reappearing (<0.5%) while I. tectus remains absent. Klausipollenites
schaubergeri dramatically declines in abundance (6.4%), Nuskoisporites dulhuntyi declines(0.5%). Protohaploxypinus sp. is rare (1.3%), with little change in the species remaining fromthe Cycle 2 carbonates; P. chaloneri (1.5%), P. jacobii (0.5%), and P. microcorpus (0.5%).
Taeniaesporites spp. overall experiences a slight increase in abundance (5.1%) and areorganization of species as T. noviaulensis appears (1.0%) while T. novimundi disappears. T.
albertae (1.3%) and T. angulistriatus (0.5%) maintain rare abundances. Striatoabieites richteri
remains absent and Striatopodocarpites sp. disappears. Vittatina hiltonensis continues to berare (<0.5%). Alisporites nuthallensis (1.0%) as well as Perisaccus granulosus (0.5%) maintainlow abundances. Falcisporites zapfei disappears. However, some species experience increases
in abundance such as Labiisporites granulatus (1.8%), Nuskoisporites cf. rotatus appears for

773
774
775
776

the first time (<0.5%), Platysaccus radialis reappears but is very rare (<0.5%), Potonieisporites
novicus increases slightly in abundance (2.6%). Reduviasporonites sp. reappears at
considerable abundance (8.4%). Cycadopites rarus, acritarchs and tetrads remain absent while
spores maintain a rare abundance (0.8%).

777
778

5.3.5. Cycle 3 Carbonates

779
780
781
782
783
784
785
786
787
788
789
790
791
792
793
794

The Cycle 3 carbonate assemblages display a continuing trend of increasingly abundant
Unidentifiable pollen grains (89.1%) at the expense of identifiable species. Lueckisporites
virkkiae is present at yet again slightly lower abundances (4.5%), however all variants are
distinguishable. Alisporites nuthallensis becomes very rare (<0.5%). Illinites sp. declines again
in abundance (3.1%), along with I. delasaucei (0.5%), while I. tectus maintains its very rareabundance (<0.5%). Klausipollenites schaubergeri continues to decline (5.9%), as does
Labiisporites granulatus (1.5%). Nuskoisporites cf. rotatus (<0.5%), N. dulhuntyi (<0.5%), andPotonieisporites novicus (0.5%) maintain rarity. No individual species of Protohaploxypinus
are distinguishable, and Protohaploxypinus sp. maintains rarity (1.0%). Reduviasporonites sp.
dramatically decreases from 8.8% to extremely rare (<0.5%). Taeniaesporites spp. experiences
a considerable decrease in abundance (1.6%) with T. labdacus (0.7%), T. noviaulensis (1.0%)
and T. angulistriatus remaining rare (1.0%). Vestigisporites minutus decreases further inabundance (3.2%). Striatoabieites richteri remains absent, Perisaccus granulosus, Platysaccus
radialis and Vittatina hiltonensis are not present. However, Striatopodocarpites antiquus
reappears but is rare (0.5%) alongside Falcisporites zapfei (<0.5%). Trisaccate pollen grainsare very rare (<0.5%).

795
796

5.3.6. Cycle 3 Evaporites

797
798
799
800
801
802
803
804
805
806
807
808
809
810
811
812
813
814
815
816
817
818

In the Cycle 3 evaporites many species experience an increase in abundance
accompanied by a decline in the proportion of Unidentifiable pollen grains (60.9%). Alisporites
nuthallensis maintains rarity (1.8%), Crustaesporites globosus reappears but is rare (<0.5%),
Cycadopites rarus occurs for the only time in the Zechstein succession (0.5%), Falcisporites
zapfei maintains rarity (0.9%). Illinites sp. increases (5.0%) and I. delasaucei (4.2%) and I.
tectus (5.2%) experience large increases, while I. klausi is absent. The abundance ofKlausipollenites schaubergeri returns to Cycle 2 levels (14.1%), and Labiisporites granulatus
also recovers (5.1%). Lueckisporites virkkiae also starts to recover but does not reach early
Zechstein abundances (8.9%), however all three variants are distinguishable, with Variant A
notably accounting for 18.6% of assemblages. Nuskoisporites dulhuntyi is still rare (0.8%) as
well as Potonieisporites novicus (0.9%), and Perisaccus granulosus reappears (0.9%). All
species of Protohaploxypinus are present with undifferentiated individuals only comprising
1.1% of assemblages. P. cf. samoilovichii (0.5%), P. chaloneri (1.5%), P. jacobii (1.7%), andP. microcorpus (1.1%) reappear. Reduviasporonites sp. increases slightly (1.8%),
Striatopodocarpites antiquus is still rare (1.2%) and Striatopodocarpites sp. reappears (1.1%).
Taeniaesporites sp. increases (2.9%) and T. albertae (2.0%) and T. novimundi (1.7%) reappear,
while T. angulistriatus maintains low abundance (1.1%). T. labdacus (2.6%) and T.
noviaulensis (1.8%) both increase marginally in abundance. Vestigisporites minutus increases
in abundance (5.9%), Vittatina hiltonensis reappears although is very rare (<0.5%). However,some species do not recover. Nuskoisporites cf. rotatus is absent. Platysaccus radialis andStriatoabieites richteri remain absent, as do tetrads. However, acritarchs (0.5%) and trisaccate
pollen grains (0.5%) were recovered although no spores were recovered.

819
820

5.3.7. Cycle 4 Carbonates

821
822
823
824
825
826
827
828
829
830
831
832
833
834
835
836
837
838
839
840
841
842

The Cycle 4 carbonate assemblages are once again dominated by Lueckisporites
virkkiae (52.0%), with all three variants distinguishable. The Unidentifiable component is
considerably reduced (13.9%). Other species also experience increases in abundance.
Crustaesporites globosus increases slightly (1.0%). Nuskoisporites cf. rotatus reappears(1.0%) however, N. dulhuntyi decreases slightly (0.5%). Platysaccus radialis reappears at veryrare abundances (<0.5%). Striatoabieites richteri reappears and Striatopodocarpites fusus
appears for the first time, both at very rare abundances (<0.5%). Taeniaesporites sp. increases
slightly (3.0%), with increases also seen in T. angulistriatus (2.3%), and T. labdacus (5.9%).
T. bilobus appears for the first and only time (0.5%). T. albertae (1.3%), T. noviaulensis (0.7%)
and T. novimundi (0.7%) maintain their rarity alongside Vittatina hiltonensis (0.7%).
Vestigisporites minutus more or less maintains abundance (5.6%). Other species experience
noticeable reductions. Klausipollenites schaubergeri declines slightly in abundance (12.1%),
Alisporites nuthallensis maintains low abundance (0.5%), while Illinites sp. reduces slightly(3.0%), with slight decreases in I. delasaucei (2.4%) and I. tectus (2.0%), while I. klausi
reappears at low abundances (0.5%). Perisaccus granulosus (0.5%) and Potonieisporites
novicus (0.5%) maintain rare abundances. Protohaploxypinus sp. (1.5%) is present at low
abundance within individual species also occurring at low abundance or being rare; P.
chaloneri (1.4%), P. jacobii (0.9%) and P. microcorpus (0.5%). Reduviasporonites sp. declines
slightly (1.8%), Striatopodocarpites antiquus disappears. Cycadopites rarus andStriatopodocarpites sp. are absent. Acritarchs maintain a rare abundance (0.5%), while tetrads
and spores remain absent. No trisaccate pollen grains are recovered after the Cycle 4
carbonates.

843
844

5.3.8. Cycle 4 evaporites and Cycle 5

845
846
847
848
849
850
851
852
853
854

The assemblages from Cycle 5 carbonates and evaporites are comparatively
impoverished with samples having low yields, a trend that appears to start in the Cycle 4
evaporites. The Cycle 4 evaporites contain an assemblage dominated by Unidentifiable pollen
grains (84.2%), accompanied by the fungus/algae Reduviasporonites sp. (22.8%),
Lueckisporites virkkiae (15.7%), Protohaploxypinus cf. samoilovichii (9.1%), andKlausipollenites schaubergeri (5.8%). In the Cycle 5 carbonates only Unidentifiable pollen
grains (96.0%) were recovered from borehole material, and benthic coiled foraminiferal test
linings (15.0%) from outcrop at Little Scar Beach (samples LSSC 2 and LSSC3). The Cycle 5
evaporites from borehole material only contain sparse Illinites sp. (5.0%), Lueckisporites
virkkiae (2.0%) and an abundance of Unidentifiable pollen grains (93.0%).

855
856

5.4. Comparison of the Durham and Yorkshire Sub-basins

857
858
859

Comparisons between spore-pollen assemblages from the Yorkshire and Durham Subbasins are only possible for the lower cycles as the upper cycles from the Durham Sub-basin
have not yielded palynomorphs.

860
861
862
863

In terms of the lowest cycle Cycle 1 The Marl Slate/‘Lower Magnesian Limestone’
assemblages in Salterford Farm borehole (Yorkshire Sub-basin) contain more taxa thanClaxheugh Rock and Crime Rigg Quarry (Durham Sub-basin). However, differences are likely
the result of differential preservation as the proportion of Unidentifiable pollen grains is

864
865
866
867
868
869
870
871
872
873
874
875
876
877

considerably lower in Salterford Farm compared to the samples from Claxheugh Rock andCrime Rigg Quarry, and in general samples from the Marl Slate of the Durham Sub-basin are
more hyaline in appearance than those from the Yorkshire Sub-basin. Illinites delasaucei,
Klausipollenites schaubergeri, Lueckisporites virkkiae Variant A, Vestigisporites minutus are
the most common species in Salterford Farm. Lueckisporites virkkiae, Taeniaesporites spp.,
Illinites spp., Vestigisporites minutus and Klausipollenites schaubergeri are most abundanttaxa in Claxheugh Rock, and Illinites spp., Klausipollenites schaubergeri, L. virkkiae,
Taeniaesporites spp. and Vestigisporites minutus are the most common species in Crime Rigg
Quarry. In addition, Taeniaesporites spp. is more abundant in the Durham Sub-basin. It appears
that Vestigisporites minutus is not a common component of early Zechstein palynofloras. V.
minutus is present in low abundance at Kimberley and in Woolsthorpe Bridge borehole, butmaintains considerable presence through the Salterford Farm borehole. In the Durham Subbasin V. minutus is only present in one sample from Claxheugh Rock yet is reasonably
abundant in the single sample from Crime Rigg Quarry.

878
879
880
881
882
883
884
885
886
887
888
889
890
891

The Cadeby Formation exposed in the Kimberley railway cutting, that was
palynologically analysed by Clarke (1965), was resampled. The assemblages described here
correspond to samples K5 and K6 of the original study. Our findings are very similar to thoseof Clarke (1965), but with some minor differences that concern observations on taxa that are
rare in the assemblages. Our study did not recover Striatopodocarpites cancellatus orLabiisporites granulatus. The reprocessed K5 sample revealed the presence of Taeniaesporites
angulistriatus, Striatopodocarpites antiquus, Vittatina hiltonensis, Illinites tectus, Perisaccus
granulosus, Nuskoisporites dulhuntyi, Potonieisporites novicus and Alisporites nuthallensis.
Both the original study and this study note the absence of Vestigisporites minutus, a species
that is abundant throughout the borehole and outcrop material. Alisporites nuthallensis was
identified in this analysis of the Kimberley material and different Protohaploxypinus species
were identified in Clarke’s study. These two genera are both rare meaning any disparity inpresence and abundance is likely due to the rarity of these two taxa and the low probability ofall species occurring in all of the slides made from the same sample.

892
893
894
895
896
897
898
899

Previous work on the Cycle 1 carbonates from Woolsthorpe Bridge (Warrington, 1980;
Berridge et al., 1999) recorded the presence of a characteristic Zechstein miospore assemblage
composed of Alisporites sp., Crustaesporites cf. globosus, Falcisporites zapfei,
Klausipollenites schaubergeri, Lueckisporites virkkiae, ?Perisaccus granulosus,
Protohaploxypinus spp., P. cf. chaloneri, P. cf. jacobii, P. microcorpus, ?Striatopodocarpites
sp., Taeniaesporites spp., T. labdacus, T. noviaulensis. Labiisporites granulatus,
Taeniaesporites albertae, and Vestigisporites minutus were not recorded (G. Warrington pers.
comm.) but have been recovered during this study.

900
901
902
903

Recovery of an assemblage from the Cadeby Formation, comparable to those from the‘Lower Marl’ at Kimberley, Cinderhill, and Woolsthorpe Bridge, has previously been reported
from Salterford Farm (Warrington, 1980; Berridge et al., 1999). The age of these assemblages
has been interpreted as Lopingian based on the presence of Lueckisporites virkkiae.

904
905

6. Reconstructing the Zechstein flora906

6.1. General comments: palaeoecology

907
908
909

The Zechstein Sea was located within the Permian Euramerican phytogeographicalprovince with a flora dominated by abundant conifers and pteridosperms with rare
ginkgophytes, sphenophytes, ferns, lycopsids and potentially cycads (Schweitzer 1986; Cleal

910
911
912
913
914
915
916
917
918
919
920
921

and Thomas, 1995). Detailed descriptions of Lopingian Euramerican flora have been provided
for England, Germany and Poland (e.g. Kurtze, 1839; Geinitz, 1869; Solms-Laubach, 1884;
Gothan and Nagelhard, 1923; Weigelt, 1928, 1930; Stoneley, 1958; Schweitzer, 1960, 1962,
1968, 1986; Ullrich, 1964; Poort and Kerp, 1990; Brandt, 1997; Uhl and Kerp, 2002), Spain(Bercovici et al., 2009), and the southern Alps (e.g. Clement-Westerhof, 1984, 1987, 1988;
Visscher, 1986; Kutstatcher et al., 2012, 2014, 2017; Labandiera et al., 2016). A Zechsteintype flora has also been reported from Belgium (Florin, 1954). The floras are generally foundin deposits of marginal marine or fluvial lowstand settings (e.g., Ullrich, 1964, Weigelt,
1928; Weigelt, 1930; Schweitzer, 1968; Schweitzer, 1986; Uhl and Kerp, 2002). It appears thatthe vegetation was fairly uniform across the basin, low in diversity and dominated by conifers(see Table 2), suggesting an arid to semi-arid environment and with plants adapted to periodic
water stress.

922
923
924
925
926
927

The British Zechstein flora has been described in detail by Stoneley (1958) andSchweitzer (1986). Due to the suggested Lopingian climate trend towards increasingaridification the Zechstein flora has been interpreted as most abundant during Cycle 1 and theprincipal transgression and gradually disappearing from Cycle 2 onwards (Schweitzer, 1986).
A similar trend of gradual decline was also suggested based on previous interpretation of
palynomorph assemblages (Pattison et al., 1973; Smith et al., 1974).

928
929
930
931
932
933

Botanically based climatic inferences can be made from anatomical structures or bycomparisons with other similar Lopingian Euramerican floras. The conifers and peltasperms
from the Zechstein Basin and southern Europe exhibit some xerophorphic adaptations. They
have very thick papillate cuticles, deeply sunken stomata, stomatal pores covered byoverarching papillae, and some conifers leaves seem to have been thick and fleshy. However,
not all taxa in the plant assemblages are fully adapted to xerophytic conditions.

934
935
936
937
938
939
940
941
942
943
944
945
946
947
948
949

The British Zechstein flora was previously divided into two groups based on theirpalaeoecology (Schweitzer, 1986). A xerophytic Callipteris-conifer association, whichcorresponds to the Callipteris-Walchia association of Gothan and Gimm (1930), and ahygrophylic Neocalamites-Sphenopterid association that corresponds to the Calamitespecopterid (fern) association of Gothan and Gimm (1930). However, given more recent
reconstructions of Lopingian Euramerican vegetation (e.g., Bercovici et al., 2009; Kutstatcher
et al., 2012, 2014; Labandiera et al., 2016; Kustatscher et al., 2017) a different structure is
proposed. It is likely that conifers (e.g., Ullmannia, Pseudovoltzia, Ortesia) with their more
xerophytic adaptations occupied the arid to semi-arid, well-drained, inland and hinterland areas
and low-lying slopes of the Protopennines. Pteridosperms (peltasperms) (e.g., Peltaspermum)
with their thicker cuticles likely inhabited coastal areas, living in drier lowland patches, and in
the hinterlands with the conifers. Horsetails (Neocalamites) likely occupied wet lowland
riparian environments, shallow water coastal bogs and lakes, and the mouth of rivers.
Pteridosperms (e.g. Sphenopteris) likely inhabited slightly less humid habitats while
ginkgophytes (e.g. Sphenobaieria) and potential cycads (e.g. Pseudoctenis, Taeniopteris)inhabited more humid lowland areas near bodies of water.

950
951
952
953
954
955
956
957

The dominance of taeniate bisaccate pollen of conifer and pteridosperm affinity in thisstudy concurs with previous investigations. While their dominance in assemblages likely
reflects the general nature of the parent vegetation, it should be noted that multitaeniate pollen
grains were produced by multiple plant groups (Chaloner, 2013), and there may also be ataphonomic bias towards bisaccate pollen due to their thick exines. Pollen grains of probableginkgo and possible cycad (Cycadopites rarus) affinity are rare. This is expected as
ginkgophytes are known to be rare components of the Zechstein flora, with some exceptions(e.g. Bauer et al. 2014), and potential cycads even rarer. However, this may also be the result

958
959

of a taphonomic bias. Finally, interpreting the exact nature and habitat of the vegetation is also
complicated by the lack of autochthonous plant remains beyond Cycle 1.

960
961

6.2. Vegetation change through time

962
963
964
965
966
967

The changing nature of spore-pollen assemblages through the Zechstein sequences ofnortheast England, as reported in this study, can be interpreted to document the changing natureof the flora. The affinities of many of the spore-pollen taxa are well documented. However, insome cases the lack of in situ occurrences means that some abundant bisaccate taxa, such as
Labiisporites granulatus, and some monosaccate pollen taxa, such as Vestigisporites minutus
and Perisaccus granulosus, have not yet been assigned to a parent plant group.

968
969
970
971
972
973
974
975
976
977

The palynological data spans almost the entire temporal extent of the Zechstein,allowing a reinterpretation of the flora. Plants were assumed to gradually disappear after Cycle
2 due to aridification and high rates of evaporation, with aridity being enhanced by thedecreasing magnitude of successive transgressive episodes. Instead, the assemblages from theupper Zechstein Cycle 4 and Cycle 5 show that the conifer-pteridosperm dominated flora
persisted through to the Permian-Triassic boundary. This was likely facilitated by increasing
fluvial activity, seen in the increase in terrigenous fluvial sediments, from the end of Cycle 3
onwards. Presumably, this provided sufficient humidity to support the flora. Thus, it appears
likely that the flora had an azonal distribution that was more severely influenced by edaphic
factors than larger scale climate change.

978
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While the upper Zechstein (Cycle 4-5) assemblages suggest a decline in the flora
towards the Permian-Triassic boundary this may also be a taphonomic effect. The assemblages
recovered from the Carnallitic Marl Formation (Cycle 4) in SM11 (Figure 10) and SM4 (Figure11) yield palynomorph assemblages of similar composition and abundance to those recoveredfrom the Marl Slate (Cycle 1) (Figure 4-6), suggesting that conditions were not as inhospitable
during the upper Zechstein as previously assumed.

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In the British Isles the Zechstein can be divided either into the classic five carbonateevaporites cycles or seven evaporite-carbonate sequences based on Tucker’s (1991) sequencestratigraphic scheme for the basin margins. The pollen charts in Figures 4-14 have beenpresented against both organisations. The sequence stratigraphic approach is particularly
appropriate as the locations studied here are marginal marine. Changing shorelines would have
had significant impacts on the Zechstein vegetation, with highstands and lowstands effecting
groundwater levels, precipitation, and therefore the distribution of suitable wetter habitats andtaphonomic windows. Interpreting the data at the scale of transgression-regression cycles orsequences may reveal the responses of vegetation to the accompanying patterns of drastic
climatic and environmental changes.

994
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1002

Long term changes across the Zechstein sequence and slow rates of change are likely areflection of climate and environmental trends of increasing aridity and temperature thatcharacterise the Lopingian. The Zechstein Group covers the last ~6 million years of thePermian meaning any slow changes may be indicative of floral turnover associated with theend-Permian mass extinction. While the assemblages of Cycle 5 are comparatively
impoverished and composed of highly degraded Unidentifiable pollen grains, this may also be
indicative of longer, water-borne transport from inland or hinterland areas via the fluvial
depositional system of Cycle 5 instead of reflecting the true nature of the state of the flora.
Furthermore, there is no coincident increase in spores with the reduction in Cycle 5

1003
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assemblages that typifies the floral turnover at the Permian-Triassic boundary. Yet, this may
also be a taphonomic effect since assemblages are generally poorly preserved.

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The cyclic transgressions into the Zechstein Basin would have forced the hydrologicalcycle by shifting coastlines and altering local topography. During highstands increased runoff
likely caused high groundwater stages in lowland areas already characterised by more
hygrophytic flora. During lowstands reduced runoff may have led to increased drainage anddesiccation of lowlands with the low-lying Protopennines already characterised by more
xerophytic flora. This can explain coeval occurrences of wet and dry lowlands and intrazonal
vegetation throughout the Zechstein. Palynological assemblages may be expected to reflect
these changing conditions, yet this is not necessarily reflected in the results presented here.

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Ginkgophytes, horsetails ferns, lycopsids and possible cycads are among the rarest
elements in the Zechstein flora, living around lakes and abandoned river channels, or along
streams in a distal flood plain setting. While this is supported by their relative absence in the
microfossil record, and may reflect the true nature of the vegetation, it may also be ataphonomic effect. Their dependence on the vicinity of large bodies of water or elevated groundwater levels would have made them more sensitive to changes in sea level accompanying theZechstein cycles.

1020
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The magnitude of sea level change diminished with each cycle as the sea progressively
shrank in volume. The hygrophytic flora would have become increasingly stressed by the
restriction of its habitat and the diminishing taphonomic window resulting in their rarity in thefossil record as both micro- and macrofossils. While their absence may be explained bysparsely distributed wet habitats suitable for reproduction, e.g., bryophytes are dependent on
moist substrate upon which to reproduce (Whitaker and Edwards, 2010), it may reflect the poor
preservation potential of certain palynomorph morphologies relative to the thicker walled
conifer and pteridosperm bisaccate pollen. For example, horsetails are known to thrive indisturbed, anoxic, and saline environments (Husby, 2013) making them ideally suited to coastal
Zechstein environments. Therefore, the rarity of horsetails in the microfossil record may be
explained by the poor preservation potential of their palynomorphs. Modern horsetails(Equisetum) produce spores that are enveloped by four flexible ribbon-like elaters that are
incredibly fragile (Marmottant et al., 2013). Furthermore, horsetails have a remarkable ability
to reproduce vegetatively via rhizomes, compensating for the inefficiency of reproduction via
spores, allowing horsetails to rapidly colonize coastal environments (Hauke, 1963), whilst
remaining under-represented in the microfossil record.

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Sea level highstands may explain the preferential preservation of coastal, nearshore,and inland habitats in certain strata, for example the Marl Slate (Cycle 1 transgression) and
Carnallitic Marl Fm. (Cycle 3-4 boundary). Both are known for an abundance of microfossil
remains, with the Marl Slate being notable for its macrofossil remains due to favourable
taphonomic conditions. Increasing energy levels during the initial Zechstein transgression
caused by the middle Wuchiapingian sea level highstand (Legler et al., 2011; Legler andSchneider, 2013) led to the rapid burial of material and the movement of azonal, coastal, flora
into the taphonomic window created by this marginal environment. The transgression alsoestablished a distal fluvial plain with meandering and abandoned channels (like those at Pot
Riding, Cadeby Formation, Cycle 1), resulting in the impression of a richer and lusher
vegetation (e.g. Kustatscher et al., 2017). The Carnallitic Marl Fm. marks the initiation of ahumid fluvial environment and therefore is also affected by a similar favourable taphonomic
bias. With higher sedimentation rates the preservation potential of the Marl Slate andCarnallitic Marl Fm. was considerably higher than that of other sedimentary settings resulting
in a higher local diversity relative to other palynomorphs assemblages.

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Unfortunately, the scarcity of plant macrofossils from beyond Cycle 1 means it is notpossible to investigate the effects of sea level change on macrofossil preservation in the upper
Zechstein, when sea level changes were of a lesser magnitude relative to the lower Zechstein,e.g., 1-2 meters rather than the sea level drop of 100-150m between Cycle 1 and Cycle 2 (Smith,1989). The absence of the Cycle 1 evaporites (Hayton Anhydrite/Hartlepool Anhydrite) means
there is no palynological data for the first major regression and evaporative phase and the biotic
responses of the flora cannot be reconstructed. By Cycle 2 the vegetation had alreadyexperienced a drastic environmental transition between arid and marine-buffered conditions
during the evaporative phase of Cycle 1.

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There are several possible trends discernible from the Zechstein palynoflora. Thereappears to be an overall reduction in palynomorph diversity and abundance through theZechstein Group, punctuated by taphonomic effects. Lueckisporites virkkiae consistently
dominates assemblages by up to ~50.0%. It is a typical Zechstein taxon recovered in the Baltic(Podoba, 1975), throughout Europe (e.g. Klaus, 1963; Clarke, 1965; Visscher, 1971; Massari
et al., 1988; Warrington and Scrivener, 1988; Massari et al., 1994, 1999; Pittau, 1999; Legler
et al., 2005; Pittau, 2005; Legler and Schneider, 2008; Warrington, 2005, 2008; Gibson et al.,2020) and from age equivalent deposits in the U.S.A. (Wilson, 1962; Clapham, 1970). It is an
important and distinct conifer signal throughout the Zechstein.

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The Marl Slate palynoflora is abundant, corroborating previous studies of taxonomic
diversity and relative abundance of palynomorph taxa, mirroring previous reports of themacrofossil record (Stoneley, 1958; Schweitzer, 1986). At the top of the Cycle 2 Fordon
Evaporite Fm. there appears to be an increase in pollen abundance seen in SM4 (Figure 11)
and SM11 (Figure 10). In particular, Unidentifiable pollen increase in abundance approaching
the Cycle 2-3 boundary. This may be in response to increasing sea level during the lateregressive-early transgressive phase expanding the distribution of coastal habitats and thetaphonomic window. This trend is also visible in SM14b (Figure 13) where Unidentifiable
pollen grains increase over the Cycle 2-3 boundary and the number of taxa present decline
across the Grauer Salzton into the Brotherton Fm. Lueckisporites virkkiae and Taeniaesporites
spp. survive the boundary, and Nuskoisporites appears. Before the boundary, an assemblage of
Illinites delasaucei, Klausipollenites schaubergeri, Labiisporites granulatus, Lueckisporites
virkkiae, Potonieisporites novicus, Reduviasporonites, Taeniaesporites sp. and Vestigisporites
minutus is present. It would appear a strong taphonomic bias is in effect as Unidentifiable
pollen grains come to utterly dominate assemblages throughout the Brotherton Fm.

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Preservation within the Cycle 3 evaporites (Billingham Anhydrite) of SM11 (Figure10) is exceptional and the intermittent recovery throughout the evaporites is related to a lower
sampling resolution through the evaporites instead of barren samples. The Carnallitic Marl Fm.
palynoflora provides unique inside into the Cycle 3-Cycle 4 boundary flora. There is a
noticeable increase in identifiable pollen species over the boundary. The assemblages
recovered from the Carnallitic Marl Fm. from SM4 and SM11 (Figures 10-11) illustrate how
conifers represent the most dominant component of the vegetation in the upper Zechstein e.g.,
Lueckisporites virkkiae, Taeniaesporites sp., and Illinites sp. Pollen grains of probably
pteridosperm affinity occur less frequently e.g. Protohaploxypinus sp., Striatoabieites sp.,
Striatopodocarpites sp. and Vittatina hiltonensis. Throughout Cycle 4 some species disappear
including Vittatina sp. (Vittatina hiltonensis) (Plate II, 1, 2). It is known to be rare in Lopingian
assemblages (Clarke, 1965; Variakhuna, 1971) yet it appears intermittently though boreholes
SM4 (Figure 11) an SM11 (Figure 10), at abundances no greater than 1.0%, until the Cycle 4
carbonates after which it disappears.

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SM11 (Figure 10) has the longest temporal range of all boreholes studied and provides
the best insight into the uppermost Zechstein flora. Palynomorphs were recovered across the
Carnallitic Marl Fm.-Sleights Siltstone-Littlebeck Anhydrite boundaries, but then tail off andonly Unidentifiable pollen grains are recovered through the rest of the sequence. The upper
Zechstein trends are heavily affected by sampling resolution and the effect of the depositionalsettings on the quality of preservation. The relative proportions of conifer and pteridosperm
pollen does not shift with conifer pollen being consistently more abundant. This trend continues
into the uppermost Zechstein, with the recovery of Illinites spp. from the Little Scarassemblages, confirming the dominance of conifers throughout the Zechstein.

1107
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The palynflora does not suggest evidence of vegetation destabilisation which is
associated with the end-Permian mass extinction (Looy et al., 2001; Lindström andMcLoughlin, 2007; Hochuli et al., 2010; Xiong and Wang, 2011; Hochuli et al., 2016;
Schneebeli-Hermann et al., 2017; Fielding et al., 2019; Novak et al., 2019). Instead it suggests
an azonal vegetation dominated by conifers and pteridosperms inhabited a semi-arid landscape
up to the Permian-Triassic boundary.

1113
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7. Palynofacies and the Zechstein palaeoenvironments

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The fossil record is a result of the complex interplay between biotic and abiotic factors,
and the composition of each palynomorph association is controlled by ecological factors, thedepositional environment and taphonomic processes. Therefore, disentangling preservationalbiases from biological signals is vital for palaeoecological reconstructions. Zechstein sporepollen assemblages are a case in point.

1120
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The Zechstein Group is interpreted as mostly marine in origin, albeit rather unusual due
to varying and increased salinity related to the evaporation-replenishment cyclicity, with theupper Zechstein (Cycles 4-5) recognised as predominantly fluvial-terrestrial. However, thepaucity of marine palynomorphs remains surprising for the lower Zechstein (Cycles 1-3). The
palynomorph assemblages are entirely dominated by allochthonous forms derived from theadjacent land mass: pollen, spores, and land plant cuticles (Stoneley, 1958). Presumably, thesewere transported in from the land by a combination of wind and water. The latter was probably
most significant during siliciclastic phases of sedimentation and surely must have been theprincipal transporter of large fragments of plant cuticle. The former may, however, have been
important in transporting pollen into hypersaline pools from which the evaporites were
deposited. Acritarchs, indicative of marine influence, are surprisingly rare. They are absent
from most samples despite sieving at 10µm and, where present, constitute no more than 0.5%of assemblages. In this study we only report Micrystridium-type from Cycle 3 evaporites andCycle 4 carbonates, although Wall and Downie (1963) previously reported them from Cycle 1
only. The lack of acritarchs presumably reflects the harsh hypersaline marginal conditions thatwere not conducive for marine phytoplankton to thrive.

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The rare foraminiferal test linings from Salterford Farm and Little Scar beach are ofinterest both for their rarity and location in the stratigraphic column, specifically for the
specimens recovered from Little Scar. These may represent the latest occurrence of Permian
foraminifera recovered in the UK (B. Spencer pers. comm.). Encrusting foraminifera have beenwell-documented from the Polish Zechstein reef systems (Peryt et al., 2012) where they form
important constituents of the limestone formations during the early Zechstein. These test
linings are benthic coiled forms, but their low quality of preservation makes taxonomic
identifications difficult. The foraminifera from Salterford Farm are not unexpected as the sea
during Marl Slate times was representative of relatively normal shallow marine conditions.

1145
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However, from Little Scar, close to the Permian-Triassic boundary, benthic foraminifera
appear in sediments interpreted as accumulating in the arid and hypersaline environments of
the later Zechstein cycles.

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Reduviasporonites in the British Permian has been correlated with brackish
environments (Warrington, 2008). This is compatible with the pattern of recovery of thesepalynomorphs in this study. This reconstruction suggests that pockets of wetter environment
persisted throughout the Zechstein. A fungal affinity has been invoked for Reduviasporonites
with it interpreted as a saprophytic agent responsible for the degradation of widespreadabundant dead organic matter preceding the end-Permian mass extinction (Visscher et al.,
1996, 2011; Hochuli, 2016). Mass occurrences of Reduviasporonites have been interpreted as
an end-Permian fungal spike, coincident with vegetation die-off and environmental disturbanceprior to and during the extinction. However, high abundances of Reduviasporonites have also
been attributed to ‘algal blooms’ (Elsik, 1999; Afonin et al., 2001; Foster et al., 2002; Spina etal., 2015). Regardless of affinity, mass occurrences of Reduviasporonites are an important
stratigraphical marker of the Permian-Triassic boundary in terrestrial and shallow marine
environments. Interestingly, Reduviasporonites maintains a rare to low abundance throughout
the Zechstein sequence, but occurs at higher abundances at the top of the Fordon Evaporite Fm.
(8.4%) (Cycle 2 (EZ3 A, K-Mg, Na) and in the Cycle 4 evaporites (Sherburn Anhydrite,
Sneaton Halite) (22.8%). However, recovery at this high abundance was only recorded in one
sample of <100 yield (see Table 1 Appendix B and Appendix E SM11 1478.30 m) which is
not conclusive evidence of a mass occurrence. The lack of a definitive “spike” in abundance atthe top of the Zechstein sequence may be viewed as further evidence that there was no
significant deforestation or vegetation turnover during Zechstein times.

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A variety in quality of preservation is observed in the Zechstein material: (i)assemblages dominated by beautifully preserved palynomorphs of low thermal maturity (e.g.
from the Marl Slate); (ii) assemblages dominated by very well-preserved palynomorphs thatare dark in colour. (e.g. from the Carnallitic Marl Fm in SM11 Dove’s Nest); (iii) assemblages
dominated by AOM but containing well-preserved yet hyaline palynomorphs which need to be
stained with Bismarck Brown to facilitate identification (e.g. from the Marl Slate at ClaxheughRock); (iv) assemblages of abundant Unidentifiable pollen grains of very poor quality
preservation; (v) Assemblages containing a mixture of types ii-iv above (e.g. argillaceous halite
samples containing well-preserved intact palynomorphs as well as highly degraded
Unidentifiable grains).

1178
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The evaporite palynomorph assemblages are of particular interest because they containa mixture of pollen preservation types, which can be explained by two preservational processes.
Well-preserved, intact palynomorphs were likely blown to the site of deposition on the wind,
settled on the surface of the super-saturated sea water where they were encapsulated in the
rapidly growing halite crystals. Poorly preserved pollen grains originate from clayey orargillaceous layers/pockets within the salt and arrived at the site of deposition via different
means, having been subjected to higher degrees of biological degradation. Some may have
washed in from the land via rivers or streams. Others may have settled out from the marine
water column. While reworking within individual halite units is a possibility due to the highmobility of halite, the possibility of reworking between evaporite units is discounted based on
the relative immobility of anhydrite and potassium-magnesium salts which constrain the moremobile layers of halite. The preservation potential of the Zechstein evaporites is discussed in
more detail in Gibson and Bodman (2020).

1191
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The dark colour of palynomorphs in some assemblages (particularly Type (ii) above)is a curious matter. Across EZ1-5 an apparent reverse thermal maturity profile indicated by

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1198

palynomorph colour is observed with older assemblages being paler and more translucent than
younger ones. Instead of being a true reverse thermal maturity profile darkening may be a resultof the thermal conductivity of evaporites in combination with the exposure of palynomorphs
to a hypersaline environment. It is also be a taphonomic effect of palynomorphs from the Marl
Slate of the Durham Sub-basin which are exceptionally hyaline in appearance, possibly due tothe more basinward location of the Durham Sub-basin.

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8. Conclusions

1201
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Euramerica during the Permian was characterised by a progressive trend towards morearid climates (Roscher and Schneider, 2006; Montañez et al., 2007). However, the semi-arid
climates of the Zechstein Basin were punctuated by several periods of more humid conditions
associated with marine transgressions.

1205
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Spore-pollen preservation potential is highest during transgressive phases and justbefore regressive phases (e.g. Kustatscher et al., 2017). Other points of cycles fall outside ofthis taphonomic window of preservation, resulting in a reduced recovery of palynomorph. The
use of evaporite palynology helps compensate for this, providing insight into the regressive
phase flora for the first time.

1210
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Pollen-dominated assemblages have been recovered from throughout the ZechsteinGroup. Assemblages are dominated by conifer pollen, reflecting the characteristics of thegymnospermous Euramerican parent flora. The composition of assemblages remains relatively
unchanged, both within and between Zechstein cycles. This confirms the reported uniform
nature of the Zechstein vegetation.

1215
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Despite the additional taphonomic window afforded by the evaporites there remains
difficulties disentangling taphonomic effects from biological signals. Regardless, thegymnospermous inland and hinterland flora appears to have persisted throughout the course of
Zechstein deposition at least until the sea finally disappeared at or close to the Permian-Triassic
boundary.

1220
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Acknowledgements.

1222
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MEG was funded by a NERC studentship through the ACCE (Adapting to the
Challenges of a Changing Environment) Doctoral Training Partnership [grant number
1807541]. We thank Geoffrey Warrington for taxonomic advice; David Bodman for assistancein the preparation of evaporite palynology samples; Asher Haynes of Sirius Minerals Plc and
the York Potash Ltd for coordinating access to boreholes SM4 Gough, SM7 Mortar Hall, SM11
Dove’s Nest and SM14b Woodsmith Mine North Shaft; Tracey Gallagher of the British
Geological Survey for coordinating access to boreholes Salterford Farm and Woolsthorpe
Bridge; the Yorkshire Wildlife Trust for approving access to the Pot Riding exposure at theSprotbrough Gorge site; Doncaster Council for approving access to the Levitt Hagg Hole site;
the owners of Transgear Units for providing access to the Sandal House exposure; Geoffrey
Clayton for discussions on reverse thermal maturity profiles; Brian Spencer for the donation of
the Little Scar Seaton Carew samples; Rick Ramsdale of the Sheffield Area Geology Trust forfieldwork assistance at the Pot Riding, Sandal House and Levitt Hagg Hole Sites. Finally, we
thank the two reviews whose comments greatly improved the manuscript.

1236

1237

SUPPORTING INFORMATION

1238

Additional Supporting Information can be found in the online version of this article:

1239

Appendix A. Lithostratigraphical and facies descriptions for all localities

1240

Appendix B. List of borehole samples, their lithology, and yield

1241

Appendix C. List of outcrop samples, their lithology, and yield

1242

Appendix D. Location and stratigraphic range of all localities

1243

Appendix E. Raw count data

1244
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References

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Afonin, S.A., Barinova, S.S., Krassilov, V.A., 2001. A bloom of Tympanicysta Balme 1980
(green algae of zygnematalean affinities) at the Permian–Triassic boundary. Geodiversitas 23,
481–487.

1249
1250

Anderson, J.M., Anderson, H.M., 1983. Palaeoflora of southern Africa, Molteno Formation(Triassic) 1. (Part 1. Introduction; Part 2. Dicroidium). Balkema, Rotterdam.

1251
1252
1253

Archangelsky, S., Wagner, R.H., 1983. Glosspteris anatolica sp. nov. from uppermost Permianstrata in south east Turkey. Bulletin of the British Museum (Natural History). Geology 37, 81–
91.

1254
1255

Balme, B.E., 1995. Fossil in situ spores and pollen grains: an annotated catalogue. Review of
Palaeobotany and Palynology 87, 81–323.

1256
1257
1258

Bauer, K., Kustatscher, E., Butzmann, R., Fischer, T.C., van Konijnburg-van Cittert, J.H.A.,
Krings, M., 2014. Ginkgophytes from the Upper Permian of the Bletterbach Gorge (Northern
Italy). Rivisita Italiana di Paleontologia e Stratigrafia 120(3), 271-279.

1259
1260

Benton, M.J., Twitchett, R.J., 2003. How to Kill (Almost) All Life: The End-Permian
Extinction Event. Trends in Ecology and Evolution 18(7), 358–365.

1261
1262
1263
1264

Bercovici, A., Diez, J.B., Broutin, J., Bourquin, S., Linol, B., Villanueva-Amadoz, U., LópezGómez, J., Durand, M., 2009. A palaeoenvironmental analysis of Permian sediments in
Minorca (Balearic Islands, Spain) with new palynological and megafloral data. Review of
Palaeobotany and Palynology 158, 14–28.

1265
1266
1267

Berridge, N.G., Pattison, J., Samuel, M.D.A., Brandon, A., Howard, A.S., Pharaoh, T. C.,
Riley, N.J., 1999. Geology of the Grantham district. Memoir of the British Geological Survey
127, 1-133.

1268
1269
1270

Bomfleur, B., Serbet, R., Taylor, E.L., Taylor, T.N. 2011. The possible pollen cone of theLate Triassic conifer Heidiphyllum/Telemachus (Voltziales) from Antarctica. Antarctic
Science 23, 379–385.

1271
1272
1273

Brauns, C.M., Pätzold, T., Haack, U., 2003. A Re-Os study bearing on the age of theKupferschiefer at Sangerhausen (Germany). International Congress on Carboniferous andPermian Stratigraphy, Utrecht, August 2003, 66 pp.

1274
1275

Chaloner, W.G., 2013. Three Palynological Puzzles. International Journal of Plant Sciences
174(3), 602-607.

1276
1277

Clarke, R.F.A., 1965. British Permian saccate and monosulcate miospores. Palaeontology 8,
322–354.

1278
1279
1280

Clayton, G. Goodhue, R., Adelbagi, S. T., Vecoli, M., 2017. Correlation of Palynomorph
Darkness Index and vitrinite reflectance in a submature Carboniferous well section in northern
Saudi Arabia. Revue de Micropaléontologie 60, 411–416 ARAMCO-CIMP.

1281
1282

Cleal, C.J., Thomas, B.A., 1995. Palaeozoic Palaeobotany of Great Britain, GeologicalConservation Review Series, No. 9, Chapman and Hall, London, 295 pp.

1283
1284
1285
1286

Clement-Westerhof, J.A., 1984. Aspects of Permian palaeobotany and palynology. IV. The
conifer Ortiseia Florin from the Val Gardena Formation of the Dolomites and the Vicentinian
Alps (Italy) with special reference to a revised concept of the Walchiaceae (Göppert) Schimper.
Review of Palaeobotany and Palynology 41, 51-166.

1287
1288
1289

Clement-Westerhof, J.A., 1987. Aspects of Permian palaeobotany and palynology. VII. The
Majonicaceae, a new family of Late Permian conifers. Review of Palaeobotany and Palynology
52, 375–402.

1290
1291
1292

Clement-Westerhof, J.A., 1988. Morphology and phylogeny of Paleozoic conifers. 298-337.
In: Beck, C. (Ed.), Origin and Evolution of Gymnosperms. Columbia University Press, New
York, 504 pp.

1293
1294

Doornenbal, J.C., Stevenson, A.G. (Eds.) 2010. Petroleum Geological Atlas of the Southern
Permian Basin Area. EAGE Publications b.v., Houten, 342 pp.

1295
1296
1297

Elsik, W.C., 1999. Reduviasporonites Wilson 1962: Synonymy of the Fungal Organism
Involved in the Late Permian Crisis. American Association of Stratigraphic Palynologists 23,
27-41.

1298
1299

Erwin, D.H., 1993. The Great Paleozoic Crisis: Life and Death in the Permian. Columbia
University Press, New York. 327 pp.

1300
1301

Erwin, D.H., 2006. Extinction: How Life on Earth Nearly Ended 250 Million Years Ago.
Princeton University Press, Princeton, New Jersey. 296 pp.

1302
1303

Florin, R., 1944. Die Koniferen des Oberkarbons und des Unteren Perms. Palaeontographica
Abteilung B-Palaophytologie 85, 365–456.

1304
1305
1306

Foster, C.B., Afonin, S.A. 2005. Abnormal pollen grains: an outcome of deteriorating
atmospheric conditions around the Permian-Triassic boundary. Journal of the GeologicalSociety of London 162, 653-659.

1307
1308
1309

Foster, C.B., Stephenson, M.H., Marshall, C., Logan, G.A., Greenwood, P. F. 2002. A Revisionof Reduviasporonites Wilson 1962: Description, Illustration, Comparison and BiologicalAffinities. American Association of Stratigraphic Palynologists 26, 35-58.

1310
1311
1312

García-veigas, J., Cendón, D.I., Pueyo, J.J., Peryt, T. M., 2011. Zechstein saline brines in
Poland, evidence of overturned anoxic ocean during the Late Permian mass extinction event.
Chemical Geology 290, 189-201.

1313
1314

Geinitz, H.B., 1869. Über fossile Pflanzenreste aus der Dyas von Val Trompia. Neues Jahrbuch
für Mineralogie, Geologie and Paliontologie 456-461.

1315
1316

Geluk, M., 1999. Late Permian (Zechstein) rifting in the Netherlands: models and implications
for petroleum geology. Petroleum Geoscience 5, 189–199.

1317
1318
1319

Gibson, M.E., Taylor, W.A., Wellman, C.H., 2020. Wall ultrastructure of the Permian pollen
grain Lueckisporites virkkiae Potonié et Klaus 1954 emend. Clarke: Evidence for botanicalaffinity. Review of Palaeobotany and Palynology 275, 104169.

1320
1321

Gibson, M.E. Bodman, D., 2020. Evaporite palynology of the British Zechstein salts: a method
for processing all evaporite types. In review.

1322
1323

Geluk, M. 2005. Stratigraphy and tectonics of Permo-Triassic basins in the Netherlands andsurrounding areas. Unpublished PhD Thesis, Utrecht University.

1324
1325

Glennie, K.W., 1983. Early Permian (Rotliegendes) Palaeowinds of the North Sea.
Sedimentary Geology 34, 245–265.

1326
1327
1328

Glennie, K.W. and Buller, A., 1983. The Permian Weissliegend of N.W. Europe: the partial
deformation of aeolian sand caused by the Zechstein transgression. Sedimentary Geology 35,
43-81.

1329
1330

Gomankov, A. V., Meyen, S. V., 1986. Tatarian flora (composition and distribution in the
Late Permian of Eurasia). Nauka, Moscow. 175 (in Russian).

1331
1332

Goodhue, R., Clayton, G., 2010. Palynomorph darkness index (PDI) - A new technique forassessing thermal maturity. Palynology 34(2), 147-156.

1333
1334

Gothan, W., Gimm, O., 1930. Neuere Beobachtungen und Betrachtungen über die Flora desRotliegenden in Thüringen. Arb Inst Palaeobot Petrograph Brennsteile 2, 39–74.

1335
1336
1337

Gotahn, W., Nagalhard, K., 1921. Kupferschieferpflanzen aus dem niederrheinischen
Zechstein. Jahrbuch der Königlich Preussischen Geologischen Landesanstalt und
Bergakademie 42, 440–460.

1338
1339

Gould, R.E., Delevoryas, T., 1977. The biology of Glossopteris: evidence from petrified seedbearing and pollen-bearing organs. Alcheringia 1, 387–399.

1340
1341
1342

Grauvogel-Stamm, L., 1978. La flore du Grès à Voltzia (Buntsandstein súperieur) des Vosges
du Nord (France). Morphologie, anatomie, interprétations phylogénique et paléogéographique.
Université Louis-Pasteur de Strasbourg Institut de Géologie 50, 1-225.

1343
1344

Grauvogel-Stamm, L., Doubinger, J., 1975. Deux fougères fertiles du Stéphanien du Massif
Central (France). Geobios, 6, 409–421.

1345
1346
1347

Grauvogel-Stamm, L., Grauvogel, L., 1973. Masculostrobus acuminatus nom. nov., un nouvel
organe reproducteur mâle de Gymnosperme de Grès à Voltzia (Trias inférieur) des Vosges(France). Geobios 6, 101–114.

1348
1349

Hallam, A., Wignall, P. B. 1997. Mass Extinctions and their Aftermath. Oxford University
Press, Oxford, 320 pp.

1350
1351
1352

Haubold, H., Schaumberg, G., 1985. Die Fossilien des Kupferschiefer: Pflanzen-und Tierwelt
zu Beginn des Zechsteins; eine Erzlagerstatte und ihre Palaontologie. Wittenberg Lutherstadt.
223 pp.

1353
1354

Hauke, R.L., 1963. A taxonomic monograph of the genus Equisetum subgenus Hippochaete.
Nova Hedwigia 8, 1-123.

1355
1356
1357

Hernandez-Castillo, G.R., Rothwell, G.W., Mapes, G., 2001. Thucydiaceae fam. nov., with areview and reevaluation of Paleozoic walchian conifers. International Journal of Plant Sciences
162, 1155–1185.

1358
1359

Hochuli, P.A., 2016. Interpretation of “fungal spikes” in Permian-Triassic boundary sections.
Global and Planetary Change 144, 48-50.

1360
1361

Hug, N., 2004. Sedimentgenese und Paläogeographie des höheren Zechsteins bis zur Basis desBuntsandsteins in der Hessischen Senke. Geologische Abhandlungen Hessen 113, 1–238.

1362
1363
1364

Hug, N. Gaupp, R., 2006. Palaeogeographic reconstruction in red beds by means of geneticallyrelated correlation: results from the upper part of the German Zechstein (Late Permian).
Zeitschrift der Deutschen Gesellschaft für Geowissenschaften 157, 107–120.

1365
1366

Husby, C., 2013. Biology and functional morphology of Equisetum with emphasis on the giant
horsetails. Botanical Review 79(2), 147-177.

1367
1368

Käding, K.C., 2000. Die Aller-, Ohre-, Friesland- und Fulda-Folge (vormals BröckelschieferFolge). Kali und Steinsalz 13, 760–770.

1369
1370

Klaus, W., 1963. Sporen aus dem südalpinen Perm. Jahrbuch der Geologischen Bundesanstalt,
Wien 106, 229–363.

1371
1372
1373

Kustatscher, E., Konijnenburg-van Cittert, J.H.A. van, Bauer, K., Butzmann, R., Meller, B.,
Fischer, T.C., 2012. A new flora from the upper Permian of Bletterbach (Dolomites, N-Italy).
Review of Palaeobotany and Palynology 182, 1–13.

1374
1375
1376
1377

Kustatscher, E., Bauer, K., Butzmann, R., Fischer, T.C., Meller, B., van Konijnenburg-van
Cittert, J.H.A., Kerp, H., 2014. Sphenophytes, pteridosperms and possible cycads from theWuchiapingian (Lopingian, Permian) of Bletterbach (Dolomites, Northern Italy). Review of
Palaeobotany and Palynology 208, 65-82.

1378
1379
1380
1381

Kustatscher, E., Bernardi, M., Petti, F.M., Franz, M., van Konijnenburg-van Cittert, J.H.A.,
Kerp, H., 2017. Sea-level changes in the Lopingian (late Permian) of the northwestern Tethys
and their effects on the terrestrial palaeoenvironments, biota and fossil preservation. Global
and Planetary Change 148, 166–180.

1382
1383

Labandiera, C.C., Kustatscher, E., Wappler, T., 2016. Floral assemblages and patterns of insect
herbivory during the Permian to Triassic of Northeastern Italy. PLoS ONE 11(11), e0165205.

1384
1385
1386

Legler, B., Gebhardt, U., Schneider, J.W., 2005. Late Permian non-marine-marine transitional
profiles in the central Southern Permian Basin, northern Germany. International Journal of
Earth Sciences 94(5–6), 851–862.

1387
1388
1389
1390

Legler, B., Schneider, J., 2008. Marine ingressions into the Middle/Late Permian saline lake of
the Southern Permian Basin (Rotliegend, Northern Germany) possibly linked to sea-levelhighstands in the Arctic rift system. Palaeogeography, Palaeoclimatology, Palaeoecology
267(1-2), 102-114.

1391
1392
1393

Lindström, S., McLoughlin, S., Drinnan, A.N., 1997. Intraspecific variation of taeniate
bisaccate pollen within Permian glossopterid sporangia, from the Prince Charles Mountains,
Antarctica. International Journal of Plant Sciences 158, 673–684.

1394
1395

Marmottant, P., Ponomarenko, A., Bienaimé, D., 2013. The walk and jump of Equisetum
spores. Proceedings of the Biological Society 280(1770), 20131465.

1396
1397
1398
1399

Menning, M., Gast, R., Hagdorn, H., Käding, K.-C., Simon, T., Szurlies, M., Nitsch, E., 2005.
Zeitskala für Perm und Trias in der Stratigraphischen Tabelle von Deutschland 2002,
zyklostratigraphische Kalibrierung der höheren Dyas und Germanischen Trias und das Alterder Stufen Roadium bis Rhaetium 2005. Newsletters on Stratigraphy 41(1/3), 173–210.

1400
1401
1402
1403
1404
1405
1406

Menning, M., Alekseev, A.S., Chuvashov, B.I., Davydov, V.I., Devuyst, F.X., Forke, H.C.,
Grunt, T.A., Hance, L., Heckel, P.H., Izokh, N.G., Jin, Y.G., Jones, P.J., Kotlyar, G.V., Kozur,
H.W., Nemyrovska, T.I., Schneider, J.W., Wang, X.D., Weddige, K., Weyer, D., Work, D.M.,
2006. Global time scale and regional stratigraphic reference scales of Central and West Europe,
East Europe, Tethys, South China, and North America as used in the Devonian–Carboniferous–
Permian Correlation Chart 2003 (DCP 2003). Palaeogeography, Palaeoclimatology,
Palaeoecology 240, 318–372.

1407
1408
1409
1410

Metcalfe, I., Foster, C.B., Afonin, S.A., Nicoll, R.S., Mundil, R., Xiaofeng, W., Lucas, S.G.
2009. Stratigraphy, biostratigraphy and C-isotopes of the Permian–Triassic non-marine
sequence at Dalongkou and Luaogou, Xinjiang Province, China. Journal of Asian Earth
Sciences 36, 503–520.

1411
1412

Meyen, S.V., 1997. Permian conifers of western Anagaraland. Review of Palaeobotany and
Palynology 96, 351–447.

1413
1414

Meyen, S.V., 1984. Basic features of gymnosperm systematics and phylogeny as shown in thefossil record. The Botanical Review 50, 1–111.

1415
1416
1417

Naugolnykh, S.V., 2014. A new genus of male cones of voltzialean affinity, Uralostrobus
voltzioides nov. gen., nov. sp., from the Lower Permian of the Urals (Russia). Geobios 47, 315324.

1418
1419
1420
1421

Massari, F., Conti, M.A., Fontana, D., Helmold, K., Mariotti, N., Neri, C., Nicosia, U., Ori,
G.G., Pasini, M., Pittau, P., 1988. The Val Gardena Sandstone and BellerophonFormation in
the Bletterbach gorge (Alto Adige, Italy): biostratigraphy and sedimentology. Memorie di
Scienze Geologiche 40, 229–273.

1422
1423
1424
1425

Massari, F., Neri, C., Pittau, P., Fontana, D., Stefani, C., 1994. Sedimentology,
palynostratigraphy and sequence stratigraphy of a continental to shallow-marine rift-related
succession: Upper Permian of the eastern Southern Alps (Italy). Mem. Sci. Geol. 46, 119–
243.

1426
1427
1428
1429
1430
1431

Massari, F., Neri, C., Fontana, D., Manni, R., Mariotti, N., Nicosia, U., Pittau, P.,
Spezzamonte, M., Stefani, C., 1999. Excursion 3: The Bletterbach section (Val Gardena
Sandstone and Bellerophon Formation), in: Stratigraphy and Facies of the Permian Deposits
between Eastern Lombardy and Western Dolomites - Field Trip Guidebook. Presented at theInternational Fiel Conference on “The Continental Permian of the Southern Alps and Sardinia(Italy). Regional Reports and General Correlations,” Brescia, Italy, pp. 111–134.

1432
1433
1434

Osborn, J.M., Taylor, T.N., 1993. Pollen morphology and ultrastructure of theCorystospermales: permineralized in situ grains from the Triassic of Antarctica. Review of
Palaeobotany and Palynology 79, 205–219.

1435
1436

Pancost, R.D., Crawford, N., Maxwell, J.R., 2002. Molecular evidence for basin-scale photic
zone euxinia in the Permian Zechstein Sea. Chemical Geology 188, 217–227.

1437
1438
1439
1440

Pattison, J., Smith, D.B., Warrington, G., 1973. A review of Late Permian and Early Triassic
biostratigraphy in the British Isles. 220-260. In: Logan, A., Howse, L.V. (eds), 1973. The
Permian and Triassic Systems and their Mutual Boundary. Memoir (Canadian Society ofPetroleum Geologists) 2, 766 pp.

1441
1442

Paul, J., 2006. Der Kupferschiefer: Lithologie, Stratigraphie, Fazies and Metallogenese eines
Schwarzschiefers. Zeitschrift der Deutschen Gesellschaft für Geowissenschaften 157, 57–76.

1443
1444
1445

Peryt, T.M., Geluk, M.C., Mathiesen, A., Paul, J., Smith, K., 2010. Zechstein. 123-147. In:
Doornenbal, J. C., Stevenson, A. G. (Eds.) Petroleum Geological Atlas of the Southern Permian
Basin Area. EAGE Publications b.v., Houten, 342 pp.

1446
1447
1448

Peryt, T. Raczyński, P., Peryt, D. and Chłódek, K., 2012. Upper Permian reef complex in thebasinal facies of the Zechstein Limestone (Ca1), western Poland. Geological Quarterly 47, 537533.

1449
1450
1451

Pittau, P., 1999. Excursion 3: the Bletterbach section (Val Gardena Sandstone andBellerophon Formation). 3. Palynology. Stratigraphy and facies of the Permian deposits
between Eastern Lombardy and the Western Dolomites. Field trip guidebook 23–25.

1452
1453
1454
1455

Pittau, P., Kerp, H., Kustatscher, E., 2005. “Let us meet across the P/T boundary”—
workshop on Permian and Triassic Palaeobotany and Palynology, Bozen. The Bletterbach
canyon. Naturmuseum Südtirol/Museo Scienze Naturali Alto Adige/Museum Natöra Südtirol,
p. 26.

1456
1457
1458

Podoba, B.G. 1975. Spores and pollen. In: Suveizdis, P. (ed.), Permian deposits of the Baltic
area (stratigraphy and gauna); Lietuvos Geologijos Mokslinio Tyrimo Institutas, Darbai,
Mintis, Vilnius 29, 184-192 (In Russian; English summary p. 196-207).

1459
1460
1461
1462

Poort, R.J., Kerp, H., 1990. Aspects of Permian palaeobotany and palynology XI. On therecognition of true peltasperms in the Upper Permian of western and central Europe and areclassification of species formerly included in Peltaspermum Harris. Review of Palaeobotany
and Palynology 63, 197-225.

1463
1464
1465
1466
1467

Poort, R. J., Clement-Westerhof, J.A., Looy, C.V., and Visscher, H., 1997. Aspects of Permian
palaeobotany and palynology XVII. Conifer extinction in Europe at the Permian-Triassic
junction: Morphology, ultrastructure and geographic/stratigraphic distribution of
Nuskoisporites dulhuntyi (prepollen of Ortiseia, Walchiaceae). Review of Palaeobotany and
Palynology 97(1-2), 9-39.

1468

Potonié, R., 1962. Synopsis der Sporae in situ. Beiheste zum Geologische Jahrbuch 52, 1–204.

1469
1470

Raine, J.I., Mildenhall, D.C., Kennedy, E.M., 2011. New Zealand Fossil Spores and Pollen:
An Illustrated Catalogue, 4th edition. GNS Science miscellaneous series no. 4.

1471
1472

Retallack, G.J., 2002. Lepidopteris callipteroides, an early Triassic seedfern of the Sydney
Basin, southeastern Australia. Alcheringa 1, 475–500.

1473
1474

Richter-Bernburg, G., 1955. Stratigraphische Gliederung des deutschen Zechstein. Zeitschrift
für Geologische Wissenschaften 105, 843–854.

1475
1476
1477

Röhling, H.-G., 1993. Der Untere Buntsandstein in Nordwest- und Nordostdeutschland – Ein
Beitrag zur Vereinheitlichung der stratigraphischen Nomenklatur. Geologisches Jahrbuch A
142, 149–183.

1478
1479

Rothwell, G. W., Mapes, G. and Hernandez-Castillo, G. R., 2005. Hanskerpia gen. nov. andphylogenetic relationships among the most ancient conifers (Voltziales). Taxon 54, 733–750.

1480
1481

Schweitzer, H.-J., 1960. Die Makroflora des niederrheinischen Zechsteins. Fortschritte
Geologisches Rheinland Westfalen 6, 1–46

1482
1483

Schweitzer, H.-J., 1968. Die Flora des Oberen Perms in Mitteleuropa. Naturwissenschaften
Rundschau 21, 93–102.

1484
1485
1486

Schweitzer, H.-J., 1986. The land flora of the English and German Zechstein sequences. In:
Harwood, G.M., Smith, D.B. (eds) The English Zechstein and Related Topics. GeologicalSociety, London, Special Publications 22, 31–54.

1487
1488
1489

Słowakiewicz, M., Kiersnowski, H., Wagner, R. 2009. Correlation of the Middle and Upper
Permian marine and terrestrial sedimentary sequences in Polish, German, and USA WesternInterior Basins with reference to global timemarkers. Palaeoworld 18, 193–211.

1490
1491
1492

Smith, D.B., Brunstrom, R.G.W., Manning, P.I., Simpson, S., Shotton, F.W., 1974. A
Correlation of the Permian Rocks of the British Isles. Geological Society, London, Special
Reports 5, 45 pp.

1493
1494

Smith, D.B., 1989. The late Permian palaeogeography of north-east England. Proceedings of
the Yorkshire Geological Society 47, 285-312.

1495
1496

Solms-Laubach, H. Graf Von, 1884. Die Coniferenformen des deutschen Kupferschiefers undZechsteins. Palaeontologische Abhandlungen 2(2), 81–116.

1497
1498
1499
1500

Spina, A., Cirilli, S., Utting, J. and Jansonius, J., 2015. Palynology of the Permian and Triassicof the Tesero and Bulla sections (Western Dolomites, Italy) and consideration about theenigmatic species Reduviasporonites chalastus. Review of Palaeobotany and Palynology 218,
3–14.

1501
1502

Stephenson, M.H., 2018. Permian palynostratigraphy: a global overview. Geological Society
Special Publication 450(1), 321-347.

1503
1504
1505

Stone, P., Milward, D., Young, B., Merritt, J.W., Clarke, S.M., McCormac, M., Lawrence,
D.J.D., 2010. British regional geology: Northern England. Fifth edition. Keworth, Nottingham:
British Geological Survey. 307 pp.

1506
1507

Stoneley, H.M.M., 1958. The Upper Permian flora of England. Bulletin of the British Museum(Natural History) Geology 3(9), 293-337.

1508
1509

Szurlies, M., 2013. Late Permian (Zechstein) magnetostratigraphy in Western and Central
Europe. Geological Society, London, Special Publication 376, 73-85.

1510

Taylor T.N., Taylor, E.L., Krings, M., 2009. Paleobotany. 2nd ed. Elsevier, Amsterdam.

1511
1512
1513

Tucker, M., 1991. Sequence stratigraphy of carbonate-evaporite basins; models and application
to the Upper Permian (Zechstein) of Northeast England and adjoining North Sea. Journal of
the Geological Society of London 148, 1019-1036.

1514
1515

Ulh, D., Kerp, H., 2002. Preservation of fossil plants from the Zechstein (Upper Permian) of
Central Europe. Freiberger Forschungshefte 497(January), 29-43.

1516
1517
1518

Variakhuna, L.M., 1971. Spores and pollen of the red beds and coal deposits of the Permianand Triassic of the northeastern region of the USSR; Akademiia Nauk SSSR, Komi Filial,
Institut Geologii, Nauka, Leningrad, 158p., pl. I-IVIII + 1-44 p. (in Russian).

1519
1520
1521

Van Wees, J.-D., Stephenson, R.A., Ziegler, P.A., Bayer, U., Mccann, T., Dadlex, R., Gaupp,R., Narkiewicz, M., Bitzer, F., Scheck, M., 2000. On the origin of the Southern Permian Basin,
Central Europe. Marine and Petroleum Geology 17, 43-59.

1522
1523

Visscher, H., 1971. The Permian and Triassic of the Kingscourt Outlier, Ireland. Special Papersof the Geological Society of Ireland 1, 1-104.

1524
1525
1526

Visscher, H., Brinkhuis, H., Dilcher, D.L., Elsik, W.C., Eshet, Y., Looy, C.V., Rampino, M.
R., Traverse, A., 1996. The terminal Paleozoic fungal event: Evidence of terrestrial ecosystem
destabilization and collapse. National Academy of Sciences Proceedings 93, 2155–2158.

1527
1528

Visscher, H., Sephton, M.A., Looy, C.V., 2011. Fungal virulence at the time of the endPermian biosphere crisis? Geology 39(9), 883-886.

1529
1530

Wagner, R., Peryt, T.M., 1997. Possibility of sequence stratigraphic subdivision of theZechstein in the Polish Basin. Geological Quarterly 41, 457–474.

1531

Wall, D., Downie, C., 1963. Permian hystrichospheres from Britain. Palaeontology 5, 770-784.

1532
1533

Warrington, G., Scrivener, R.C. ,1988. Late Permian fossils from Devon: regional geologicalimplications. Proceedings of the Usher Society 7, 95-96.

1534
1535
1536

Warrington, G. 2005. The chronology of the Permian and Triassic of Devon and south-east
Cornwall (UK): a review of methods and results. Geoscience in south-west England 11, 117122.

1537
1538

Warrington, G., 2008. Palynology of the Permian succession in the Hilton Borehole, Vale ofEden, Cumbria, UK. Proceedings of the Yorkshire Geological Society 57(2), 123-130.

1539
1540
1541

Weigelt, J., 1928. Die Pflanzenreste des mitteldeutschen Kupferschiefers und ihre Einschaltung
ins Sediment – Eine palökologische Studie. Fortschritte der Geologie und Paläontologie, VI
19, 395–592.

1542
1543

Whitaker, D.L., Edwards, J., 2010. Sphagnum moss disperses spores with vortex rings. Science
329, 406.

1544
1545

Zavada, M.S., Crepet, W.L., 1985. Pollen wall ultrastructure of the type material of Pteruchus
africanus, P. dubius and P. papillatus. Pollen Spores 27, 271–276.

1546
1547
1548

Zavialova, V., Nosova, N., Gavrilova, O., 2016. Pollen grains associated with gymnospermous
mesofossils from the Jurassic of Uzbekistan. Review of Palaeobotany and Palynology 233,
125-145.

1549
1550

Zavada, M.S., 1991. The ultrastructure of pollen found in dispersed sporangia of Arberiella(Glossopteridaceae). Botanical Gazette 152, 248–255.

1551
1552
1553

Ziegler, A.M., 1990. Phytogeographic patterns and continental configurations during thePermian period. Palaeozoic Palaeogeography and Biogeography, McKerrow, W.S., Scotese,
C.R. (Eds.), Memoirs of the Geological Society 12, 363-379.

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Plate I

1562

2

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3

6

5

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.
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.

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_

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~

. ,,.
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.

11

14

12

13

15

16

17

18

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Plate II

1563

3

2

4

7

10

8

11

12

15

16

,...l

'"

~

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~

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18

17

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'.

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.

14

13

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20

Plate III

1564
1565

3

4

7
6

8

9

11

1566

Plate Descriptions

1567
1568
1569
1570

Plate I. Taeniate bisaccate pollen grains. England Finder co-ordinates included. Scale bar
represents 50 µm in all images. Images taken using a QImaging (Model No. 01-MP3.3-RTVR-CLR-10) camera mounted on an Olympus BH-2 transmitted light microscope in conjunction
with QCapture Pro software.

1571
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1573
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1575
1576
1577
1578
1579
1580
1581
1582
1583
1584
1585
1586
1587
1588
1589
1590
1591
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1593
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1597
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1599
1600
1601
1602
1603
1604
1605
1606
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1608
1609
1610
1611
1612
1613

1. Lueckisporites virkkiae Variant A (KIM 5, P43)
2. L. virkkiae Variant A (SM11 1312.24 m, U42/1)
3. L. virkkiae Variant B (KIM 5, B33/1)
4. L. virkkiae Variant C (KIM 6, E36/1)
5. L. virkkiae Variant C (SM11 1312.24 m, K30/1)
6. Taeniaesporites labdacus (SM11 1312.24 m, W45/2)
7. T. labdacus (SM11 1312.24 m, J32/2)
8. T. noviaulensis (SM11 1312.24 m, R42)
9. T. noviaulensis (SM11 1312.24 m, M48/2)
10. T. novimundi (SM11 1312.24 m, S31/2)
11. T. angulistriatus (SM1 1312.24 m, V35/3)
12. T. albertae (SM11 1458.37 m, J46/3)
13. Protohaploxypinus chaloneri (SM11 1312.24 m, G30)
14. P. chaloneri (SM11 1312.24 m, R36/3)
15. P. jacobii (SM11 1312.24 m, S46)
16. P. jacobii (SM11 1312.24 m, L33/1))
17. P. microcorpus (SM11 1312.24 m, K33)
18. P. cf. samoilovichii. (SM11 1334.10 m, O39)
Plate II. Taeniate and non-taeniate bisaccate pollen and monosaccate pollen. . England Finder
co-ordinates included. Scale bar represents 50 µm for all images. Images taken using a
QImaging (Model No. 01-MP3.3-RTV-R-CLR-10) camera mounted on an Olympus BH-2
transmitted light microscope in conjunction with QCapture Pro software.
1. Vittatina hiltonensis (SM11 1312.24 m, F31)
2. V. hiltonensis (M4 1438.07 m, H32/2)
3. Striatoabieites richteri (SM11 1312.24 m, K27/3)
4. Striatopodocarpites antiquus (SM11 1312.24 m, P27)
5. S. fusus (SM11 1312.24 m, M29)
6. Platysaccus radialis (SM11 1312.24 m, J33/4)
7. Klausipollenites schaubergeri (SM11 1312.24 m, T39/3)
8. K. schaubergeri (SM11 1312.24 m, D32/3)
9. Vestigisporites minutus (KIM 5, P41)
10. V. minutus (SM11 1312.24 m, L30)
11. V. minutus (SM11 1312.24 m, J44/2)
12. Illinites delasaucei (SM11 1312.24 m, V41)
13. I. klausi (SM11 1328.76 m, P31/4)
14. I. tectus (SM11 1312.24 m, K30)
15. I. tectus (SM11 1312.24 m, Q41)
16. Falcisporites zapfei (SM11 1312.24 m, H42)
17. F. zapfei (SM11 1312.24 m, M33/2)
18. Alisporites nuthallensis (SM11 1312.24 m, M45)
19. Labiisporites granulatus (SM11 1312.24 m, L41)
20. L. granulatus (SM11 1312.24 m, G48/3)

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1615
1616
1617
1618
1619

21. Alisporites nuthallensis (SM11 1312.24 m, M45)
Plate III. Monosaccate pollen grains, trilete spores and foraminiferal test linings. England
Finder co-ordinates included. Scale bar represents 50 µm for all images. Images taken using a
QImaging (Model No. 01-MP3.3-RTV-R-CLR-10) camera mounted on an Olympus BH-2
transmitted light microscope in conjunction with QCapture Pro software.
1. Nuskoisporites cf. rotatus (SM11 1328.76 m, E43)
2. N. dulhuntyi (SM11 1312.24 m, T35/3)
3. Perisaccus granulosus (SM11 1312.24 m, W35/3)
4. P. granulosus (SM11 1312.24 m, O43/1)
5. Potonieisporites novicus (SM11 1304.14 m, E31)
6. ?Potonieisporites novicus (1312.24 m, D32/2)
7. Crustaesporites globosus (SM11 1328.76 m, R40/1)
8. Trilete spore (SM11 1465.92 m, P39)
9. Trilete spore (SM11 1465.92 m, C31/4)
10. Foraminiferal test lining (LSSC3, C37/4)
11. Foraminiferal test lining (SFYFP6373, S48)

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1621
1622
1623
1624
1625
1626
1627
1628
1629
1630
1631
1632
1633

Table 1. List of the taxa encountered during this study and their probable botanical affinities.

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Table 2. Summary of the distribution of the taxa encountered during this study.

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