Systematic descriptions
Division Dinoflagellata (Bütschli, 1885) Fensome et al.,
1993.
Subdivision Dinokaryota Fensome et al., 1993.
Class Dinophyceae Pascher, 1914.
Subclass Peridiniphycidae Fensome et al., 1993.
Order Gonyaulacales Taylor, 1980.
Suborder Ceratiineae Fensome et al., 1993.
Family Ceratiaceae Willey and Hickson, 1909.
Genus Odontochitina Deflandre, 1937.
Type. Odontochitina operculata (Wetzel, 1933)
Deflandre and Cookson, 1955.
Odontochitina diducta Pearce, 2010 emend. nov.
(Pl. 1, figs. 1–3)
1967 Odontochitina costata Alberti, 1961;
Clarke and Verdier: pl. 13, fig. 4 only.
1968 Odontochitina sp. Cookson and Eisenack: 112,
pl. 2, fig. D.
1991 Odontochitina operculata (Wetzel, 1933)
Deflandre and Cookson, 1955; Heine: pl. 27, fig. 15.
1992 Odontochitina sp. Costa and Davey: pl. 3.13,
fig. 5.
1997 Odontochitina operculata (Wetzel,
1933) Deflandre and Cookson, 1955; Roncaglia and Corradini: pl. 2,
fig. 5.Emended diagnosis. A cornucavate to hypocavate species of Odontochitina
with a widely divergent antapical and lateral horn separated by an angle
equal to or greater than 80∘. Emended description. Large ceratioid, cavate
dinoflagellate cyst with one apical, antapical and lateral horn of comparable
length. The wall is two-layered, comprising a smooth endophragm and smooth,
incompletely and faintly striate or distally perforate (although this may be
due to corrosion) periphragm. The periphragm and endophragm are appressed in
the precingular region with the lateral and antapical horns being either
cornucavate or connected by a hypocavation. The endocyst is sub-spherical,
lacking obvious projections into the pericoel. The antapical and lateral
horns are separated by an angle of greater than 80∘. The paracingulum
may be indicated by faint ridges on the periphragm. The archaeopyle is apical
Type (tA) and the operculum is detached. Remarks. In the original description of O. diducta, Pearce (2010:62) described the species as having “a well-developed
cavation connecting the lateral and antapical horns”. However, this feature seems to be gradational to specimens from the same material that are cornucavate, and the diagnosis and description are emended here to account for
this fact. The specimens referred to as Odontochitina sp. by Cookson
and Eisenack (1968) and Costa and Davey (1992) are thus now considered
synonymous with O. diducta.
Suborder Gonyaulacineae (autonym).
Family Areoligeraceae Evitt, 1963.
Genus Glaphyrocysta Stover and Evitt, 1978.
Type. Glaphyrocysta retiintexta (Cookson, 1965) Stover and
Evitt, 1978.
Glaphyrocysta coniacia sp. nov.
(Pl. 1, figs. 4–11)
2003 Glaphyrocysta sp. A Pearce et al.: pl. 1,
fig. 8.
Derivation of name. Named after the Coniacian Stage from
which the type material was obtained.
Diagnosis. A species of Glaphyrocysta with a
microreticulate endophragm and a weakly fibrous periphragm forming processes
of highly variable width, united into linear, arcuate, soleate or annulate
complexes that unite distally into wide, perforate, recurved and irregular
membranes.
Holotype. MPK 14626, EF coordinates: M40, Pl. 1,
figs. 4–5.
Paratypes. Paratype 1, MPK 14627, EF coordinates: K46/1,
Pl. 1, fig. 6; paratype 2, MPK 14628, EF coordinates: Q21/2, Pl. 1, fig. 7;
paratype 3, MPK 14629, EF coordinates: K19, Pl. 1, fig. 8; paratype 4, MPK
14630, EF coordinates: J11, Pl. 1, fig. 9; paratype 5, MPK 14631, EF
coordinates: L22/1, Pl. 1, figs. 10–11.
Type locality and horizon. BGS Banterwick Barn
borehole, Berkshire; 1.48–1.55 m, Upper Chalk Formation (Broadstairs Chalk
Member), Micraster coranguinum Zone (Middle Coniacian).
Description. Medium to large chorate dinoflagellate cyst
with a lenticular, dorso-ventrally compressed, central body and an offset
sulcal notch. Two weak antapical bulges may be present. Wall two-layered,
comprising a micro-reticulate endophragm and a slightly fibrous periphragm
that separate in the formation of processes. The processes are tabular,
apparently solid, and are well developed around the ambitus of the central
body, poorly developed on the mid-dorsal area and typically absent on the
mid-ventral area. The marginal processes are of variable width but similar
length on individual specimens and appear to be clustered into linear groups
that are connected distally and may be connected at the base. The mid-dorsal
processes may develop into shorter arcuate, soleate or annulate complexes.
Distally, the processes widen into perforated platforms that are irregular in
shape and typically recurved. The sulcal notch is offset to the left; the
cingulum is represented by slender processes at margins of the central body.
The archaeopyle is apical, Type tA, typically with a detached operculum.
Dimensions. Holotype, central body
w/l = 74 × 61 µm, maximum process
length = 35 µm; paratype 1, central body
w/l = 71 × 62 µm, maximum process
length = 27 µm; paratype 2, central body
w/l = 59 × 53 µm, maximum process
length = 30 µm; paratype 3, central body
w/l = 60 × 60 µm, maximum process
length = 30 µm; paratype 4, central body
w/l = 70 × 53 µm, maximum process
length = 25 µm; paratype 5, maximum process
length = 36 µm. Range, central body
w/l = 46(60)74 × 42(55)64 µm, maximum process
length = 12(29)36 µm. Sixteen specimens measured.
Stratigraphic distribution of the new species from the BGS
Banterwick Barn and Trunch boreholes, calibrated by the isotopic events of
Jarvis et al. (2006). Abbreviations: A, Albian;
Aj, Acanthoceras jukesbrownei;
BC, Ballard Cliff Member; Ce, Cenomanian;
Cg, Calycoceras guerangeri;
CTBE, Cenomanian–Turonian Boundary Event; CR, Chalk Rock;
G. quadrata, Gonioteuthis quadrata;
L, lower; LCE III, Lower Cenomanian Event III; M, middle;
M. coranguinum, Micraster coranguinum;
Mc, Micraster cortestudinarium; MCE
I, Middle Cenomanian Event I; MCE II, Middle Cenomanian Event II;
Md, Mantelliceras dixoni;
Mg, Metoicoceras geslinianum;
Ml, Mytiloides labiatus;
Mm, Mantelliceras mantelli;
Mt, Marsupites testudinarius;
Nj, Neocardioceras juddii;
Op, Offaster pilula;
Sp, Sternotaxis plana; St M's Chalk, St
Margaret's Chalk; Tl, Terebratulina lata;
U, upper; Us, Uintacrinus socialis.
Modified from Jarvis et al. (2006).
Stratigraphic range. Upper Chalk Formation, Broadstairs
Chalk Member, Micraster coranguinum Zone (Middle Coniacian), above
the White Fall Event in the Banterwick Barn borehole to an unconfirmed upper
limit (Fig. 2).
(1–3) Spiniferites
ramosus subsp. ginakrogiae subsp. nov. (holotype), MPK
14639, EF coordinates: T46: (1) internal (reversed) dorsal view;
(2) ambital view; (3) external ventral view.
(4) Spiniferites ramosus subsp.
ginakrogiae subsp. nov. (paratype 1), MPK 14640, EF coordinates:
V19/2: external (right) lateral view. (5) Spiniferites
ramosus subsp. ginakrogiae subsp. nov. (paratype 2), MPK
14641, EF coordinates: V26/4: external (left) lateral view. (6, 8) Spiniferites ramosus subsp. ginakrogiae subsp.
nov., specimen from the Gina Krog Field, Norwegian North Sea;
(7) Spiniferites ramosus subsp. ginakrogiae subsp.
nov. (left) and S. ramosus subsp. ramosus (right), size
comparison. Scale bar represents 10 µm.
Comparison. The closest species to Glaphyrocysta coniacia sp. nov. are G. exuberans (Deflandre and Cookson, 1955 ex
Eaton, 1976) Stover and Evitt, 1978, G. intricata (Eaton, 1971) Stover
and Evitt, 1978 and G. texta (Bujak, 1976) Stover and Evitt, 1978.
Despite being described from the Palaeogene and therefore being significantly
younger, Glaphyrocysta exuberans differs with processes
that ramify elaborately halfway along their length, while G. intricata and G. texta show much less variability in process width.
Glaphyrocysta intricata also differs in having processes that
bifurcate distally to a variable length and width.
Family Gonyaulacaceae Lindemann, 1928.
Subfamily Gonyaulacoideae (autonym).
Genus Spiniferites Mantell, 1850.
Type. Spiniferites ramosus (Ehrenberg, 1837) Mantell, 1854.
Spiniferites ramosus subsp.
ginakrogiae subsp. nov.
(Pl. 2, figs. 1–6, 7 (left), 8)
Derivation of name. After the Norwegian Gina Krog and oil
field (in the Norwegian North Sea) of the same name, where the species has
also been recognised.
Diagnosis. A large subspecies of Spiniferites ramosus with a smooth wall, narrow sutural crests and intergonal processes
on some of the larger plate boundaries.
Holotype. MPK 14639, EF coordinates: T46, Pl. 2,
figs. 1–3.
Paratypes. Paratype 1, MPK 14640, EF coordinates: V19/2,
Pl. 2, fig. 4; paratype 2, MPK 14641, EF coordinates: V26/4, Pl. 2, fig. 3.
Type locality and horizon. BGS Trunch borehole, Norfolk;
451.0 m, Upper Chalk (Burnham Chalk Formation), Sternotaxis
plana Zone (Upper Turonian).
Description. Large, spiniferate, chorate dinoflagellate
cyst with a sub-spherical central body. The wall is two-layered with a thick
(∼ 2 µm) and smooth endophragm and a thin
(∼ 0.5 µm) periphragm, the latter of which develops solid
processes. Distally trifurcating gonal and bifurcating intergonal processes
(on the boundaries between the larger precingular, cingular and postcingular
plates) are typically less than one-third the central body diameter, with the
furcation occurring from 50 % (on shorter processes) to 25 % (on
longer processes) from the distal end. Narrow sutural crests line the
processes, often to the distal extremity, occasionally rendering the bi- and
trifurcations relatively wide, and define a clear standard gonyaulacacean
tabulation. The cingulum is weakly laevorotatory, typically by one cingulum
width; the parasulcus lacks clearly developed sutures. The archaeopyle is
precingular, Type P3′′, operculum detached.
Dimensions. Holotype: central body w/l:
74 × 75 µm, maximum process length = 22 µm,
paratype 1: central body w/l: 71 × 76 µm, maximum process
length = 29 µm, paratype 2: central body w/l:
91 × 80 µm, maximum process length = 19 µm.
Range central body
w/l = 54(69)91 µm × 50(69)84 µm, maximum
process length, 15(21)30 µm. Twenty specimens measured.
Stratigraphic range. Ferriby Chalk Formation,
Mantelliceras mantellii Zone (Lower Cenomanian; questionably between
the Lower Cenomanian Event I and the Virgatus Beds Event) to the
Burnham–Flamborough Chalk Formation (undifferentiated), Gonioteuthis quadrata Zone (Lower Campanian) in the Trunch borehole (Fig. 2).
(1–3) Conosphaeridium norfolkense sp.
nov. (holotype), MPK 14624, EF coordinates: K37/1: internal (reversed) view
of the precingular archaeopyle; precise orientation not determined.
(4–6) Conosphaeridium norfolkense sp. nov.
(paratype), MPK 14625, EF coordinates: V26/1: external view; precise
orientation not determined. Scale bar represents 10 µm.
Remarks. This species appears to be morphologically
intermediate between Spiniferites ramosus subsp. gracilis
(by possessing intergonal processes) and Spiniferites
porosus (in size). The central body diameter of Spiniferites porosus ranges from 66 to 69 µm (the holotype is
66 × 75 µm), with process lengths ranging from
17 to 36 µm (holotype 17–23 µm) long.
Comparison. Differs from other subspecies of
Spiniferites ramosus by its conspicuously large size.
Subfamily Leptodinioideae Fensome et al., 1993.
Genus Conosphaeridium Cookson and Eisenack, 1969.
Type. Conosphaeridium striatoconum (Deflandre and
Cookson, 1955) Cookson and Eisenack, 1969.
Conosphaeridium norfolkense sp. nov.
(Pl. 3, figs. 1–6)
1976 Conosphaeridium cf. striatoconum of
Benson: pl. 1, figs. 4–5.
Derivation of name. Named after the English county of
Norfolk, from where the type material was obtained.
Diagnosis. A species of Conosphaeridium with a
smooth to only very weakly striate periphragm.
Holotype. MPK 14624, EF coordinates: K37/1, Pl. 3,
figs. 1–3.
Paratype. MPK 14625, EF coordinates: V26/1, Pl. 3,
figs. 4–6.
Type locality and horizon. BGS Trunch borehole, Norfolk;
423.0 m, Upper Chalk (Burnham Chalk Formation), Micraster
cortestudinarium Zone (Lower Coniacian).
Description. Medium-sized proximochorate dinoflagellate
with a globular sub-spherical central body. The wall is two-layered,
comprised of a thin, smooth endophragm and smooth to very weakly striate
periphragm that forms hollow intratabular processes. The processes are
relatively squat, rounded-lagenate and typically open distally. The
archaeopyle is precingular, Type P3′′, the operculum may be attached,
but more usually detached.
Dimensions. Holotype, central body
diameter = 60 µm, maximum processes
length = 17 µm. Paratype, central body
diameter = 51 µm, maximum processes
length = 12 µm. Range, central body
diameter = 44(54)65 µm, maximum processes
length = 8(12)17 µm. Six specimens measured.
Stratigraphic range. Only observed from the sample
containing the holotype, between the Beeding and Light Point events (Fig. 2).
Comparison. Both Conosphaeridium
abbreviatum Wilson, 1984 and C. striatoconum (Deflandre and
Cookson, 1955) Cookson and Eisenack, 1969 differ by possessing strong ribs on
the processes; the former also has a much larger central body size (holotype
body diameter 95 × 81 µm). Brideaux and McIntyre (1975,
pl. 7, figs. 17–18) figured a specimen as Conosphaeridium sp.
A from the Middle Albian of northern Canada that is similar to C. norfolkense in possessing a very weakly striate periphragm, but which
differs in the shape of the processes that are tubular and narrow distally
with a truncated distal margin. The specimen figured as
Conosphaeridium cf. striatoconus by Benson (1976) from the
undifferentiated Upper Cretaceous of Maryland, USA, is considered here
synonymous with C. norfolkense sp. nov.
Subfamily uncertain
Genus Sentusidinium Sarjeant and Stover, 1978.
Type. Sentusidinium rioultii (Sarjeant, 1968) Sarjeant and
Stover, 1978.
Remarks. As stated by Wood et al. (2016) in their
emendation of the genus, the operculum of Sentusidinium is normally
detached (implying that attached operculae are permitted), and deep accessory
archaeopyle sutures are typically present.
Sentusidinium devonense sp. nov.
(Pl. 4, figs. 1–12)
(1–3) Sentusidinium
devonense sp. nov. (holotype), MPK 14634, EF coordinates: O17:
(1) external view; (3) internal (reversed) view.
(4–6) Sentusidinium devonense sp. nov. (paratype 1), MPK
14635, EF coordinates: F37/3: (4) external view;
(6) internal (reversed) view. (7–9) Sentusidinium devonense sp. nov. (paratype 2), MPK 14636, EF coordinates: D50/4:
(7) external view; (9) internal (reversed) view.
(10–12) Sentusidinium devonense sp. nov.
(paratype 3), MPK 14637, EF coordinates: L16: (10) external view;
(12) internal (reversed) view. Scale bar represents
10 µm.
1987 Sentusidinium sp. B in Tocher and Jarvis:
pl. 9.3, figs. 7–9.
1995 Canningia sp. B FitzPatrick: fig. 9n.
Derivation of name. From the English county of Devon, from
where the species was originally recorded.
Diagnosis. A species of Sentusidinium with an even
covering of short, flexuous and acuminate spines.
Holotype. MPK 14634, EF coordinates: O17, Pl. 4,
figs. 1–3.
Paratypes. Paratype 1, MPK 14635, EF coordinates: F37/3,
Pl. 4, figs. 4–6; paratype 2, MPK 14636, EF coordinates: D50/4, Pl. 4,
figs. 7–9; paratype 3, MPK 14637, EF coordinates: L16, Pl. 4, figs. 10–12.
Type locality and horizon. BGS Banterwick Barn borehole,
Berkshire; 48.06–48.09 m, Middle Chalk Formation, New Pit Chalk Member,
Terebratulina lata Zone (Middle Coniacian).
Description. A medium-sized proximate dinoflagellate cyst
with a sub-rounded body. The wall is composed of a finely granular autophragm
that possesses short (∼ 2 µm), evenly distributed and
apparently solid, simple flexuous spines with acuminate tips. No expression
of the cingulum or sulcus is present. The archaeopyle is apical, Type tA,
with a zig-zag margin and clear accessory sutures. The operculum may be
attached but more usually detached.
Dimensions. Holotype, central body w/l (excluding
operculum) = 66 × 43 µm, maximum process length
2 µm; paratype 1, central body w/l (excluding
operculum) = 65 × 48 µm, maximum process length
2 µm; paratype 2, central body
w/l = 68 × 50 µm, maximum process length
2 µm; paratype 3, central body
w/l = 60 × 50 µm, maximum process length
3 µm. Range, central body w/l (excluding
operculum) = 40(61)76 × 40(48)60 µm, maximum process
length = 2(2)3 µm. Seventeen specimens measured.
Stratigraphic range. Middle Chalk Formation, Holywell Chalk
Member, Mytiloides labiatus/Terebratulina
lata boundary (Middle Turonian; between the Lulworth and Round Down
events in the Banterwick Barn borehole) to the Burnham–Flamborough Chalk
Formation (undifferentiated), Micraster cortestudinarium
Zone (Lower Coniacian; between the Navigation and Beeding events in the
Trunch borehole; Fig. 2).
Remarks. FitzPatrick (1995) described the informal species
Canningia sp. B from the Turonian of Southern England. She
considered the generic attribution based on the subspherical outline,
non-tabular spinose ornament and apical archaeopyle; however, she also
mentioned the lack of an offset sulcal notch. These observations suggest that
the species is not an areoligeracean, and that it has a greater affinity with
Sentusidinium, and it is here considered synonymous with S.
devonense sp. nov.
Comparison. For such a simple genus of dinoflagellate cyst,
there are a number of comparable species, but which all differ in the shape
of the central body and/or the morphology of the processes.
Sentusidinium aptiense (Burger, 1980a) Burger, 1980b possesses hollow
tubular spines and S. capillatum (Davey, 1975) Lentin and Williams,
1981, S. echinatum (Gitmez and Sarjeant, 1972) Sarjeant and Stover,
1978 and S? millepiedii (Jain and Millepied,
1975) Islam, 1993 possesses much more densely distributed ornament.
Sentusidinium capitatum (Cookson and Eisenack, 1960) Wood et al.,
2016 may possess short spines with acuminate tips but the body is distinctly
elongate, while S. minus (Jiabo, 1978) He et al. in He et al., 1989
is significantly smaller. Sentusidinium myriatrichum Fensome, 1979
possesses significantly shorter and denser ornament and S. perforoconum (Yun Hyesu, 1981) Islam, 1993 has a densely perforate periphragm.
Sentusidinium pilosum (Ehrenberg, 1854) Sarjeant and Stover, 1978 has
a denser cover of short processes with variable tips that are also variable
in S. rioultii Sarjeant, 1968, S. sahii (Khanna and Singh,
1981) Wood et al., 2016, S. sparsibarbatum Erkmen and Sarjeant, 1980
and S. villersense (Sarjeant, 1968) Sarjeant and Stover, 1978.
Sentusidinium separatum (McIntyre and Brideaux, 1980) Lentin and
Williams, 1981 has bifid and branched process tips.
Sentusidinium spinosum sp. nov.
(Pl. 5, figs. 1–6)
Derivation of name. From the Latin spinosum,
meaning spiny.
Diagnosis. A species of Sentusidinium with an even
covering of relatively long, flexuous and acuminate spines.
Holotype. MPK 14638, EF coordinates: S13/4, Pl. 5,
figs. 1–6.
Type locality and horizon. BGS Banterwick Barn borehole,
Berkshire; 39.55–39.59 m, Upper Chalk Formation, St. Margaret's Chalk
Member, Sternotaxis plana Zone (Middle Turonian).
Description. A medium-sized proximate dinoflagellate cyst
with a sub-rounded body. The wall is composed of a micro-reticulate
autophragm that possesses relatively long (4–16 µm), evenly
distributed, hollow spines with simple acuminate tips. No expression of the
cingulum or sulcus is present. The archaeopyle is apical, Type tA, with a
zig-zag margin and clear accessory sutures. The operculum may be attached but
more usually detached.
(1–6) Sentusidinium
spinosum sp. nov. (holotype), MPK 14638, EF coordinates: S13/4:
(1) internal (reversed) view, showing the archaeopyle margin;
(6) external view of the antapical region.
(7–9) Impletosphaeridium banterwickense sp. nov.
(holotype), MPK 14632, EF coordinates: Q35/3: (7) external view;
(9) internal (reversed) view.
(9–12) Impletosphaeridium banterwickense sp. nov.
(paratype), MPK 14633, EF coordinates: K6/3: (10) external view;
(12) internal (reversed) view. Scale bar represents
10 µm.
Dimensions. Holotype, central body
w/l = 61 × 48 µm, maximum process
length = 11 µm. Range, central body
w/l = 46(56)72 × 30(46)56 µm, maximum processes
length = 4(6)16 µm. Twenty specimens measured.
Stratigraphic range. Middle Chalk Formation, Holywell Chalk
Member, Mytiloides labiatus/Terebratulina
lata boundary (Middle Turonian; between the Lulworth and Round Down
events) to the Upper Chalk Formation, Broadstairs Chalk Member,
Micraster coranguinum Zone (Middle Coniacian; below the
White Fall Event) in the Banterwick Barn borehole (Fig. 2).
Comparison. Sentusidinium aptiense
(Burger, 1980a) Burger, 1980b also possesses hollow spines but which differ
from S. spinosum sp. nov. in their relatively shorter length of
3–4 µm. These species also have mutually exclusive ranges, being
Callovian to Aptian for S. aptiense (Wood et al., 2015) and
Turonian for S. spinosum sp. nov. Sentusidinium
densicomatum (Maier, 1959) Sarjeant, 1983 differs in possessing more
densely distributed hair-like projections. The processes are also much more
numerous on S. pilosum (Ehrenberg, 1854) Sarjeant and Stover, 1978
and which also have variable tips. Spine length is comparable on S. sahii (Khanna and Singh, 1981) Wood et al., 2016 and S. seperatum
(McIntyre and Brideaux, 1980) Lentin and Williams, 1981 but which also have
variable tips in the former and bifid and branched tips in the latter.
Suborder uncertain
Family uncertain
Genus Impletosphaeridium Morgenroth, 1966.
Type. Impletosphaeridium transfodum Morgenroth, 1966.
Impletosphaeridium banterwickense sp. nov.
(Pl. 5, figs. 7–12)
Derivation of name. Named after Banterwick Barn, the
borehole from where the species is described. Diagnosis. A species of Impletosphaeridium
possessing spines that terminate distally into a simple bifurcation with
endings of equal length. Holotype. MPK 14632, EF coordinates: Q35/2, Pl. 5,
figs. 7–9. Paratype. MPK
14633, EF coordinates: K6/3, Pl. 5, figs. 10–12.
Type locality and horizon. BGS Banterwick Barn borehole, Berkshire; 79.51–79.53 m, Middle Chalk
Formation, New Pit Chalk Member, Terebratulina lata Zone
(Middle Turonian). Description. A small chorate dinoflagellate cyst with a
sub-rounded body. The wall is composed of a finely granular autophragm from
which arises evenly distributed, non-tabular solid spines, which terminate in
a short bifurcation. No expression of the cingulum or sulcus is present. The
archaeopyle is apical, Type tA, with a zig-zag margin and accessory sutures.
The operculum may be attached but more usually detached.
Dimensions. Holotype,
central body w/l = 46 × 39 µm, maximum process
length = 12 µm. Paratype, central body
w/l = 46 × 39 µm, maximum process
length = 14 µm. Range, central body
w/l = 35(41)47 × 29(36)43 µm, maximum process
length = 10(13)16 µm. Twenty specimens measured.
Stratigraphic range. Middle
Chalk Formation, Ballard Cliff Member (uppermost Cenomanian; upper
Cenomanian–Turonian Boundary Event) in the Banterwick Barn borehole to the
Burnham–Flamborough Chalk Formation (undifferentiated), Micraster
coranguinum Zone (Middle Coniacian; between the White Fall and
Kingsdown events) in the Trunch borehole (Fig. 2).
Comparison.
Impletosphaeridium furcillatum (Prössl, 1990 ex
Prössl, 1992) Williams et al., 1998 differs in possessing thicker
bifurcate to multifurate processes, while I. ligospinosum (de
Coninck, 1969) Islam, 1983 differs in possessing bifurcations of unequal
length. In I. varispinosum (Sarjeant, 1959) Islam, 1993, the spines
are more numerous and differ in being occasionally simple but more
frequently bifurcate, clavate or “hammer-headed”. The only similar species
of comparable age is I. williamsii (Boltenhagen, 1977) Islam, 1993; although it possesses acuminate to bifurcate processes, many of the latter
style more closely resemble flared process endings than bifurcating spines. Species
of Downiesphaeridium differ in possessing hollow (and distally
closed) processes, the closest species of which is D. aciculare
(Davey, 1969) Islam, 1993, which differs further by possessing wider (although
relatively thin), blade-like processes. According to Davey (1969), the
processes of Downiesphaeridium aciculare are always pointed
distally and occasionally bear small subsidiary spines near their
extremities. In I. banterwickense sp. nov., the subsidiary spines
occur at the extremity and are significantly longer.