Geological Survey of Denmark and Greenland Bulletin 36
GEOLOGICAL SURVEY OF DENMARK AND GREENLAND BULLETIN 36· 2016
Cretaceous and Cenozoic dinoflagellate
cysts and other palynomorphs from the
western and eastern margins of the
Labrador–Baffin Seaway
Robert A. Fensome, Henrik Nøhr-Han sen
& Graham L. Williams
GEOLOGICAL SURVEY OF DENMARK AND GREENLAND
MINISTRY OF ENERGY, UTILITIES AND CLIMATE
Bulletin36.qxp_Bulletin 36 19/12/16 13.39 Side 1
Geological Survey of Denmark and Greenland Bulletin 36
Keywords
Biostratigraphy, Cretaceous, Cenozoic, dinocysts, systematics, Baffin Margin, Labrador Margin, Mesozoic, offshore West Greenland
Cover illustration
Selected palynomorphs from the subsurface of the Labrador–Baffin Seaway; for details, see Plates 1, 3, 4, 6, 7, 12, 15, 17, 18.
Top left to bottom right: Dinoflagellates – Chiropteridium gilbertii sp. nov., Chatangiella tripartite, Cleistosphaeridium palmatum sp. nov.,
Diphyes brevispinum, Ginginodinium? flexidentatum sp. nov., Adnatosphaeridium vittatum, Piladinium columna, Thalassiphora pelagica,
Eocladopyxis peniculata; Miospores – Aquilapollenites quadrilobus, Cicatricososporites eocenicus, Baculatisporites crenulatus sp. nov.
Frontispiece: facing page
The Canadian icebreaker Amundsen off Beechey Island in the Canadian Arctic in the autumn of 2013. Geological research in the Arctic is
entirely dependent on such professional logistic support, whether by sea or air. Photo: Kate Jarrett (GSC).
Chief editor of this series: Adam A. Garde
Scientific editor of this volume: Jon R. Ineson
Editorial secretary: Jane Holst
Referees: Martin Pearce (UK) and James B. Riding (UK)
Illustrators: Jette Halskov (GEUS) and Bill MacMillan (GSCA)
Digital photographic work: Benny M. Schark
Graphic production: Annabeth Andersen
Printers: Rosendahls · Schultz Grafisk a/s, Albertslund, Denmark
Manuscript received: 21 January 2015
Final version approved: 17 March 2016
Printed: 19 December 2016
ISSN (print) 1604-8156
ISSN (online) 1904-4666
ISBN (print) 978-87-7871-443-5
ISBN (online) 978-87-7871-444-2
Citation of the name of this series
It is recommended that the name of this series is cited in full, viz. Geological Survey of Denmark and Greenland Bulletin.
If abbreviation of this volume is necessary, the following form is suggested: Geol. Surv. Den. Green. Bulletin 36, 143 pp.
Available from
Geological Survey of Denmark and Greenland (GEUS)
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Phone: +45 38 14 20 00, fax: +45 38 14 20 50, e-mail: geus@geus.dk
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© De Nationale Geologiske Undersøgelser for Danmark og Grønland (GEUS), 2016
For the full text of the GEUS copyright clause, please refer to www.geus.dk/publications/bull
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Bulletin36.qxp_Bulletin 36 19/12/16 13.39 Side 3
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Contents
Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Methodology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Systematics – general . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Systematics – dinoflagellates . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Division Dinoflagellata
Class Dinophyceae
Genus Achilleodinium Eaton 1976 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Achilleodinium biformoides (Eisenack 1954) Eaton 1976
Genus Adnatosphaeridium Williams & Downie 1966a . . . . . . . . . . . . . . . . . . . . . . . .
Adnatosphaeridium vittatum Williams & Downie 1966a
Genus Alisocysta Stover & Evitt 1978 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Alisocysta circumtabulata (Drugg 1967) Stover & Evitt 1978
Alisocysta margarita (Harland 1979a) Harland 1979a
Genus Alterbidinium Lentin & Williams 1985 emend. nov. . . . . . . . . . . . . . . . . . . . .
Alterbidinium acutulum (Wilson 1967a) Lentin & Williams 1985
Alterbidinium biaperturum (McIntyre 1975) comb. nov.
Alterbidinium? bicellulum (Islam 1983a) Lentin & Williams 1985
Alterbidinium ioannidesii Pearce 2010
Alterbidinium varium Kirsch 1991
Genus Apectodinium (Costa & Downie 1976) Lentin & Williams 1977a emend. Williams,
Damassa, Fensome & Guerstein in Fensome et al. 2009 . . . . . . . . . . . . . . . . . . .
Apectodinium homomorphum (Deflandre & Cookson 1955) Lentin & Williams 1977a
Apectodinium parvum (Alberti 1961) Lentin & Williams 1977a
Apectodinium quinquelatum (Williams & Downie 1966b) Costa & Downie 1979
Genus Aptea Eisenack 1958 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Aptea polymorpha Eisenack 1958
Genus Apteodinium Eisenack 1958 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Apteodinium australiense (Deflandre & Cookson 1955) Williams 1978
Apteodinium spiridoides Benedek 1972
Genus Areoligera Lejeune-Carpentier 1938 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Areoligera circumsenonensis Fensome et al. 2009
Areoligera gippingensis Jolley 1992
Genus Areosphaeridium Eaton 1971 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Areosphaeridium diktyoplokum (Klumpp 1953) Eaton 1971
Genus Atopodinium Drugg 1978 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Atopodinium cf. haromense Thomas & Cox 1988
Genus Axiodinium Williams et al. in Fensome et al. 2009 . . . . . . . . . . . . . . . . . . . . . .
Axiodinium augustum (Harland 1979b) Williams et al. 2015
Genus Batiacasphaera Drugg 1970 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Batiacasphaera micropapillata Stover 1977
Genus Batioladinium Brideaux 1975 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Batioladinium jaegeri (Alberti 1961) Brideaux 1975
Genus Callaiosphaeridium Davey & Williams 1966a . . . . . . . . . . . . . . . . . . . . . . . . . .
Callaiosphaeridium asymmetricum (Deflandre & Courteville 1939)
Davey & Williams 1966a
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Genus Cannosphaeropsis Wetzel 1933a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Cannosphaeropsis passio de Verteuil & Norris 1996
Genus Cerebrocysta Bujak in Bujak et al. 1980 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Cerebrocysta bartonensis Bujak in Bujak et al. 1980
Cerebrocysta magna Bujak 1994 .
Genus Cerodinium Vozzhennikova 1963 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Cerodinium diebelii (Alberti 1959) Lentin & Williams 1987
Cerodinium glabrum (Gocht 1969) Fensome et al. 2009
Cerodinium kangiliense Nøhr-Hansen & Heilmann-Clausen 2001
Cerodinium speciosum (Alberti 1959) Lentin & Williams 1987
Cerodinium striatum (Drugg 1967) Lentin & Williams 1987
Genus Charlesdowniea Lentin & Vozzhennikova 1989 emend. Williams et al. 2015 . . .
Charlesdowniea coleothrypta (Williams & Downie 1966b)
Lentin & Vozzhennikova 1989
Genus Chatangiella Vozzhennikova 1967 emend. nov. . . . . . . . . . . . . . . . . . . . . . . . .
Chatangiella decorosa (McIntyre 1975) Lentin & Williams 1976
Chatangiella madura Lentin & Williams 1976
Chatangiella tripartita (Cookson & Eisenack 1960a) Lentin & Williams 1976
Genus Chiropteridium Gocht 1960 emend. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Chiropteridium galea (Maier 1959) Sarjeant 1983
Chiropteridium gilbertii sp. nov.
Genus Chlamydophorella Cookson & Eisenack 1958 . . . . . . . . . . . . . . . . . . . . . . . . . .
Chlamydophorella nyei Cookson & Eisenack 1958
Chlamydophorella cf. nyei Cookson & Eisenack 1958
Genus Chytroeisphaeridia (Sarjeant 1962) Downie & Sarjeant 1965 . . . . . . . . . . . . . .
Chytroeisphaeridia hadra sp. nov.
Genus Cleistosphaeridium Davey et al. 1966 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Cleistosphaeridium diversispinosum Davey et al. 1966
Cleistosphaeridium elegantulum sp. nov.
Cleistosphaeridium palmatum sp. nov.
Cleistosphaeridium polypetellum (Islam 1983b) Stover & Williams 1995
Genus Cordosphaeridium Eisenack 1963a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Cordosphaeridium cantharellus (Brosius 1963) Gocht 1969
Cordosphaeridium delimurum Fensome et al. 2009
Cordosphaeridium fibrospinosum Davey & Williams 1966a
Cordosphaeridium funiculatum Morgenroth 1966a
Cordosphaeridium gracile (Eisenack 1954) Davey & Williams 1966a
Cordosphaeridium inodes (Klumpp 1953) Eisenack 1963a
Genus Cribroperidinium Neale & Sarjeant 1962 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Cribroperidinium giuseppei (Morgenroth 1966a) Helenes 1984
Genus Cyclonephelium Deflandre & Cookson 1955 . . . . . . . . . . . . . . . . . . . . . . . . . . .
Cyclonephelium distinctum (Deflandre & Cookson 1955) Jansonius 1986
Genus Dapsilidinium Bujak et al. 1980 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Dapsilidinium pseudocolligerum (Stover 1977) Bujak et al. 1980
Dapsilidinium pseudoinsertum sp. nov.
Dapsilidinium simplex (White 1842) Bujak et al. 1980
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Genus Deflandrea Eisenack 1938 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Deflandrea borealis sp. nov.
Deflandrea denticulata Alberti 1959
Deflandrea galeata (Lejeune-Carpentier 1942) Lentin & Williams 1973
Deflandrea majae (Schiøler 1993) comb. nov.
Deflandrea oebisfeldensis Alberti 1959
Deflandrea phosphoritica Eisenack 1938
Genus Dinogymnium Evitt et al. 1967 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Dinogymnium longicorne (Vozzhennikova 1967) Harland 1973
Genus Diphyes Cookson 1965 nom. cons. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Diphyes brevispinum Bujak 1994
Diphyes colligerum (Deflandre & Cookson 1955) Cookson 1965
Diphyes ficusoides Islam 1983a
Genus Disphaerogena Wetzel 1933a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Disphaerogena carposphaeropsis Wetzel 1933a
Genus Eatonicysta Stover & Evitt 1978 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Eatonicysta furensis (Heilmann-Clausen in Heilmann-Clausen & Costa 1989)
Stover & Williams 1995
Eatonicysta ursulae (Morgenroth 1966a) Stover & Evitt 1978
Genus Enneadocysta Stover & Williams 1995 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Enneadocysta magna Fensome et al. 2007
Genus Eocladopyxis Morgenroth 1966a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Eocladopyxis peniculata Morgenroth 1966a
Genus Evittosphaerula Manum 1979 emend. Damassa 1997 . . . . . . . . . . . . . . . . . . . .
Evittosphaerula? foraminosa sp. nov.
Genus Fibrocysta Stover & Evitt 1978 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Fibrocysta bipolaris (Cookson & Eisenack 1965a) Stover & Evitt 1978
Genus Gillinia Cookson & Eisenack 1960a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Gillinia hymenophora Cookson & Eisenack 1960a
Genus Ginginodinium Cookson & Eisenack 1960a . . . . . . . . . . . . . . . . . . . . . . . . . . .
Ginginodinium? flexidentatum sp. nov.
Genus Glaphyrocysta Stover & Evitt 1978 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Glaphyrocysta divaricata (Williams & Downie 1966a) Stover & Evitt 1978
Glaphyrocysta exuberans (Deflandre & Cookson 1955 ex Eaton 1976)
Stover & Evitt 1978
Glaphyrocysta retiintexta Cookson 1965
Glaphyrocysta texta (Bujak 1976) Stover & Evitt 1978
Glaphyrocysta vicina (Eaton 1976) Stover & Evitt 1978
Genus Habibacysta Head et al. 1989 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Habibacysta tectata Head et al. 1989
Genus Hapsocysta Davey 1979 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Hapsocysta? benteae Nøhr-Hansen 1993
Genus Heteraulacacysta Drugg & Loeblich Jr. 1967 . . . . . . . . . . . . . . . . . . . . . . . . . . .
Heteraulacacysta porosa Bujak in Bujak et al. 1980
Genus Heterosphaeridium Cookson & Eisenack 1968 . . . . . . . . . . . . . . . . . . . . . . . . . .
Heterosphaeridium bellii Radmacher et al. 2014
Heterosphaeridium difficile (Manum & Cookson 1964) Ioannides 1986
Genus Histiocysta Davey 1969a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Histiocysta palla Davey 1969a
Genus Homotryblium Davey & Williams 1966a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
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Homotryblium abbreviatum Eaton 1976
Homotryblium tenuispinosum Davey & Williams 1966a
Genus Hystrichokolpoma Klumpp 1953 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Hystrichokolpoma cinctum Klumpp 1953
Hystrichokolpoma globulus Michoux 1985
Genus Hystrichosphaeridium Deflandre 1937 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Hystrichosphaeridium quadratum sp. nov.
Hystrichosphaeridium tubiferum (Ehrenberg 1838) Deflandre 1937
Genus Hystrichosphaeropsis Deflandre 1935 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Hystrichosphaeropsis perforata Schiøler 1993
Hystrichosphaeropsis quasicribrata (Wetzel 1961) Gocht 1976
Genus Hystrichostrogylon Agelopoulos 1964 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Hystrichostrogylon digitus sp. nov.
Genus Impagidinium Stover & Evitt 1978 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Impagidinium victorianum (Cookson & Eisenack 1965b) Stover & Evitt 1978
Genus Impletosphaeridium Morgenroth 1966a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Impletosphaeridium apodastum sp. nov.
Genus Isabelidinium Lentin & Williams 1977b . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Isabelidinium cooksoniae (Alberti 1959) Lentin & Williams 1977b
Isabelidinium cretaceum (Cookson 1956) Lentin & Williams 1977b
Isabelidinium microarmum (McIntyre 1975) Lentin & Williams 1977b
Genus Kiokansium Stover & Evitt 1978 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Kiokansium williamsii Singh 1983
Genus Kleithriasphaeridium Davey 1974 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Kleithriasphaeridium mantellii (Davey & Williams 1966a) comb. nov.
Genus Laciniadinium McIntyre 1975 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Laciniadinium arcticum (Manum & Cookson 1964) Lentin & Williams 1980
Genus Lentinia Bujak in Bujak et al. 1980 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Lentinia serrata Bujak in Bujak et al. 1980
Genus Licracysta Fensome et al. 2007 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Licracysta corymbus Fensome et al. 2007
Licracysta? semicirculata (Morgenroth 1966b) Fensome et al. 2007
Genus Lingulodinium Wall 1967 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Lingulodinium funginum (Morgenroth 1966a) Islam 1983a
Lingulodinium machaerophorum (Deflandre & Cookson 1955) Wall 1967
Genus Nyktericysta Bint 1986 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Nyktericysta davisii Bint 1986
Nyktericysta dictyophora He Chengquan et al. 1992
Nyktericysta tripenta (Bint 1986) Fensome et al. 2009
Genus Odontochitina Deflandre 1937 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Odontochitina ancala Bint 1986
Odontochitina costata Alberti 1961
Odontochitina porifera Cookson 1956
Genus Oligosphaeridium Davey & Williams 1966a . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Oligosphaeridium albertense (Pocock 1962) Davey & Williams 1969
Oligosphaeridium pulcherrimum (Deflandre & Cookson 1955)
Davey & Williams 1966a
Oligosphaeridium totum Brideaux 1971
Genus Operculodinium Wall 1967 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Operculodinium centrocarpum (Deflandre & Cookson 1955) Wall 1967
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Genus Palaeocystodinium Alberti 1961 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Palaeocystodinium bulliforme Ioannides 1986
Palaeocystodinium golzowense Alberti 1961
Palaeocystodinium teespinosum Fensome et al. 2009
Genus Palaeohystrichophora Deflandre 1935 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Palaeohystrichophora infusorioides Deflandre 1935
Genus Palaeoperidinium Deflandre 1934 ex Sarjeant 1967 . . . . . . . . . . . . . . . . . . . . . .
Palaeoperidinium pyrophorum (Ehrenberg 1838 ex Wetzel 1933b) Sarjeant 1967
Genus Palynodinium Gocht 1970 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Palynodinium grallator Gocht 1970
Genus Petalodinium Williams et al. 2015 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Petalodinium condylos (Williams & Downie 1966b) Williams et al. 2015
Genus Phelodinium Stover & Evitt 1978 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Phelodinium kozlowskii (Górka 1963) Lindgren 1984
Genus Phthanoperidinium Drugg & Loeblich Jr. 1967 . . . . . . . . . . . . . . . . . . . . . . . .
Phthanoperidinium coreoides (Benedek 1972) Lentin & Williams 1976
Phthanoperidinium levimurum Bujak in Bujak et al. 1980
Phthanoperidinium multispinum Bujak in Bujak et al. 1980
Phthanoperidinium regale Bujak 1994
Phthanoperidinium stockmansii (de Coninck 1975) Lentin & Williams 1977a
Genus Piladinium Williams et al. 2015 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Piladinium columna (Michoux 1988) Williams et al. 2015
Piladinium edwardsii (Wilson 1967b) Williams et al. 2015
Genus Pseudoceratium Gocht 1957 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Pseudoceratium sp.
Genus Raphidodinium Deflandre 1936 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Raphidodinium fucatum Deflandre 1936
Genus Reticulatosphaera Matsuoka 1983 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Reticulatosphaera actinocoronata (Benedek 1972) Bujak & Matsuoka 1986
Genus Rhombodinium Gocht 1955 emend. Williams et al. in Fensome et al. 2009 . . . .
Rhombodinium draco Gocht 1955
Rhombodinium porosum Bujak 1979
Genus Rottnestia Cookson & Eisenack 1961a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Rottnestia borussica (Eisenack 1954) Cookson & Eisenack 1961a
Genus Scalenodinium gen. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Scalenodinium scalenum sp. nov.
Genus Schematophora Deflandre & Cookson 1955 . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Schematophora speciosa Deflandre & Cookson 1955
Genus Senegalinium Jain & Millepied 1973 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Senegalinium iterlaaense Nøhr-Hansen & Heilmann-Clausen 2001
Genus Senoniasphaera Clarke & Verdier 1967 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Senoniasphaera inornata (Drugg 1970) Stover & Evitt 1978
Senoniasphaera microreticulata Brideaux & McIntyre 1975
Senoniasphaera rotundata Clarke & Verdier 1967
Genus Simplicidinium gen. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Simplicidinium insolitum (Eaton 1976) comb. nov.
Genus Sophismatia Williams et al. 2015 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Sophismatia tenuivirgula (Williams & Downie 1966b) Williams et al. 2015
Genus Spinidinium Cookson & Eisenack 1962 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Spinidinium echinoideum (Cookson & Eisenack 1960a) Lentin & Williams 1976
8
59
59
59
60
60
61
61
62
63
63
63
64
64
64
65
65
65
66
67
67
Bulletin36.qxp_Bulletin 36 19/12/16 13.39 Side 8
Genus Spiniferites Mantell 1850 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Spiniferites ovatus Matsuoka 1983
Spiniferites pseudofurcatus (Klumpp 1953) Sarjeant 1970
Spiniferites scabrosus (Clarke & Verdier 1967) Lentin & Williams 1975
Genus Spongodinium Deflandre 1936 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Spongodinium delitiense (Ehrenberg 1838) Deflandre 1936
Spongodinium grossum (Manum & Cookson 1964) comb. nov.
Spongodinium obscurum (Manum & Cookson 1964) comb. nov.
Genus Stichodinium Williams et al. 2015 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Stichodinium lineidentatum (Deflandre & Cookson 1955) Williams et al. 2015
Genus Subtilisphaera Jain & Millepied 1973 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Subtilisphaera perlucida (Alberti 1959) Jain & Millepied 1973
Genus Surculosphaeridium Davey et al. 1966 emend. nov. . . . . . . . . . . . . . . . . . . . . . .
Surculosphaeridium convocatum sp. nov.
Genus Talladinium Williams, Damassa, Fensome & Guerstein in Fensome et al. 2009 . .
Talladinium? clathratum (Eisenack 1938) Williams, Damassa, Fensome & Guerstein
in Fensome et al. 2009
Talladinium pellis sp. nov.
Genus Tanyosphaeridium Davey & Williams 1966a . . . . . . . . . . . . . . . . . . . . . . . . . . .
Tanyosphaeridium xanthiopyxides (Wetzel 1933a ex Deflandre 1937)
Stover & Evitt 1978
Genus Taurodinium gen. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Taurodinium granulatum sp. nov.
Genus Tenua Eisenack 1958 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Tenua hystrix Eisenack 1958
Genus Thalassiphora Eisenack & Gocht 1960 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Thalassiphora delicata Williams & Downie 1966a
Thalassiphora fenestrata Liengjarern et al. 1980
Thalassiphora pelagica (Eisenack 1954) Eisenack & Gocht 1960
Genus Trichodinium Eisenack & Cookson 1960 . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Trichodinium castanea Deflandre 1935 ex Clarke & Verdier 1967
Genus Trithyrodinium Drugg 1967 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Trithyrodinium? conservatum sp. nov.
Trithyrodinium evittii Drugg 1967
Trithyrodinium quinqueangulare Marheinecke 1992
Trithyrodinium suspectum (Manum & Cookson 1964) Davey 1969b
Genus Tuberculodinium Wall 1967 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Tuberculodinium vancampoae (Rossignol 1962) Wall 1967
Genus Vesperopsis Bint 1986 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Vesperopsis longicornis (Batten & Lister 1988) Harding 1990
Genus Wallodinium Loeblich Jr. & Loeblich III 1968 . . . . . . . . . . . . . . . . . . . . . . . . .
Wallodinium luna (Cookson & Eisenack 1960a) Lentin & Williams 1973
Genus Wetzeliella Eisenack 1938 emend. Williams et al. in Fensome et al. 2009 . . . . .
Wetzeliella articulata Wetzel in Eisenack 1938 emend. Williams et al. in
Fensome et al. 2009
Genus Xenascus Cookson & Eisenack 1969 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Xenascus ceratioides (Deflandre 1937) Lentin & Williams 1973
Xenascus wetzelii Slimani 1996 ex Slimani 2001a
Systematics – acritarchs and other algae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Fromea (Cookson & Eisenack 1958) Yun Hyesu 1981 . . . . . . . . . . . . . . . . . . .
9
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68
69
69
70
71
71
72
73
73
73
74
75
75
76
76
76
77
77
Bulletin36.qxp_Bulletin 36 19/12/16 13.39 Side 9
Fromea nicosia Jansonius 1989
Fromea quadrangularis sp. nov.
Genus Microsphaeridium Benedek 1972 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Microsphaeridium ancistroides Benedek 1972
Genus Palambages Wetzel 1961 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Palambages spp.
Genus Paralecaniella Cookson & Eisenack 1970a . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Paralecaniella indentata (Deflandre & Cookson 1955) Cookson & Eisenack 1970a
Genus Pediastrum Meyen 1829 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Pediastrum spp.
Genus Tetraporina Naumova 1950 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Tetraporina sp. A
Tetraporina sp. B
Systematics – miospores and fungal elements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Miospores . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus Afropollis Doyle et al. 1982 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Afropollis sp.
Genus Appendicisporites Weyland & Krieger 1953 . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Appendicisporites potomacensis Brenner 1963
Appendicisporites unicus (Markova in Ivanova & Markova 1961) Singh 1964
Genus Aquilapollenites Rouse 1957 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Aquilapollenites quadrilobus Rouse 1957 emend. Braman 2013
Genus Azolla Lamarck in Lamarck et al. 1783 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Azolla spp.
Genus Baculatisporites Pflug & Thomson in Thomson & Pflug 1953 . . . . . . . . . . . . .
Baculatisporites crenulatus sp. nov.
Genus Callialasporites Dev 1961 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Callialasporites dampieri (Balme 1957) Dev 1961
Callialasporites obrutus Norris 1969
Genus Caryapollenites Raatz 1938 ex Potonié 1960 . . . . . . . . . . . . . . . . . . . . . . . . . . .
Caryapollenites inelegans Nichols & Ott 1978
Caryapollenites veripites (Wilson & Webster 1946) Nichols & Ott 1978
Genus Cerebropollenites Nilsson 1958 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Cerebropollenites mesozoicus (Couper 1958) Nilsson 1958
Genus Chenopodipollis Krutzsch 1966 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Chenopodipollis sp.
Genus Cicatricosisporites Potonié & Gelletich 1933 . . . . . . . . . . . . . . . . . . . . . . . . . .
Cicatricosisporites minutaestriatus (Bolkhovitina 1961) Pocock 1964
Cicatricosisporites ornatus Srivastava 1972
Genus Cicatricososporites Pflug & Thomson in Thomson & Pflug 1953 . . . . . . . . . . .
Cicatricososporites eocenicus (Selling 1944) Jansonius & Hills 1976
Genus Compositoipollenites Potonié 1951 ex Potonié 1960 . . . . . . . . . . . . . . . . . . . . . .
Compositoipollenites sp. B of Williams & Brideaux 1975
Genus Corsinipollenites Nakoman 1965 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Corsinipollenites oculusnoctis (Thiergart 1940) Nakoman 1965
Genus Extratriporopollenites Pflug in Thomson & Pflug 1952 ex Pflug in
Thomson & Pflug 1953 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Extratriporopollenites spp.
Genus Graminidites Cookson 1947 ex Potonié 1960 . . . . . . . . . . . . . . . . . . . . . . . . . .
Graminidites sp. A. of Williams & Brideaux 1975
10
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80
80
80
80
80
81
81
82
82
83
83
83
83
84
84
84
84
Bulletin36.qxp_Bulletin 36 19/12/16 13.39 Side 10
Genus Momipites Wodehouse 1933 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Momipites annellus Nichols & Ott 1978
Momipites coryloides Wodehouse 1933
Genus Osmudacidites Couper 1953 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Osmundacidites wellmannii Couper 1953
Genus Parviprojectus Mtchedlishvili in Samoilovitch & Mtchedlishvili 1961emend.
Braman 2013 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Parviprojectus reticulatus Mtchedlishvili in Samoilovitch & Mtchedlishvili 1961
Genus Parvisaccites Couper 1958 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Parvisaccites amplus Brenner 1963
Parvisaccites radiatus Couper 1958
Genus Periporopollenites Pflug & Thomson in Thomson & Pflug 1953 . . . . . . . . . . . .
Periporopollenites sp.
Genus Pistillipollenites Rouse 1962 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Pistillipollenites macgregorii Rouse 1962
Genus Quercoidites Potonié et al. 1950 ex Potonié 1960 . . . . . . . . . . . . . . . . . . . . . . .
Quercoidites sp.
Genus Rugubivesiculites Pierce 1961 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Rugubivesiculites spp.
Genus Tiliaepollenites Potonié 1931 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Tiliaepollenites crassipites (Wodehouse 1933) comb. nov.
Tiliaepollenites sp. A
Genus Translucentipollis Khlonova 1961 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Translucentipollis contiguus (Tschudy 1969) Braman 2013
Genus Wodehouseia Stanley 1961 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Wodehouseia spinata Stanley 1961
Genus Zlivisporis Pacltová 1961 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Zlivisporis spp.
Genus Zonalapollenites Pflug in Thomson & Pflug 1953 . . . . . . . . . . . . . . . . . . . . . .
Zonalapollenites igniculus (Potonié 1931) Thomson & Pflug 1953
Fungal elements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Plates 1–20 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
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Bulletin36.qxp_Bulletin 36 19/12/16 13.39 Side 12
13
Abstract
Authors’ addresses
R.A.F. & G.L.W., Geological Survey of Canada (Atlantic), Natural Resources Canada, PO Box 1006, 1 Challenger Drive,
Dartmouth, Nova Scotia B2Y 4A2, Canada. E-mail: rob.fensome@canada.ca
H.N.-H., Geological Survey of Denmark and Greenland, Øster Voldgade 10, DK-1350 Copenhagen K, Denmark.
New palynological analysis of samples from 13 offshore wells on the Canadian Margin and six wells on
the West Greenland Margin has led to a new event biostratigraphic framework for Cretaceous–
Cenozoic strata of the Labrador Sea – Davis Strait – Baffin Bay (Labrador–Baffin Seaway) region.
This framework is based on about 150 dinoflagellate cyst taxa and 30 acritarch, algal, fungal and
plant microfossil (mostly miospore) taxa. In the systematics we include three new genera of
dinocysts (Scalenodinium, Simplicidinium and Taurodinium), 16 new species of dinocysts (Chirop -
teridium gilbertii, Chytroeisphaeridia hadra, Cleistosphaeridium elegantulum, Cleistosphaeridium
palmatum, Dapsilidinium pseudoinsertum, De flan drea borealis, Evittosphaerula? foraminosa, Gingi -
nodinium? flexidentatum, Hystricho sphae ridium quadratum, Hystrichostrogylon digitus, Impleto sphaer -
idium apodastum, Scalenodinium scalenum, Surculo sphaeridium convocatum, Talladinium pellis, Tau-
rodinium granulatum and Trithyrodinium? conservatum), four emendations of dinocyst genera (Alter -
bidinium, Cha tangiella, Chiropteridium and Surculosphae ridium), six new combinations for dinocyst
species (Alterbidinium biaperturum, De flandrea majae, Kleithriasphaeridium mantellii, Simplicidinium
insolitum, Spongodinium grossum, Spon godinium obscurum), one new acritarch species (Fromea quadran-
gularis), one new miospore species (Bacula ti s porites crenulatus) and one new combination for miospores
(Tiliaepollenites crassipites). Most of the taxa included provide age information, almost exclusively last
occurrences (range ‘tops’), but some are useful mainly for environmental interpretations. Collectively,
they provide a powerful tool for helping to establish the geological history of the Labrador–Baffin Seaway.
Fensome, R.A., Nøhr-Hansen, H., & Williams. G.L. 2016: Cretaceous and Cenozoic
dinoflagellate cysts and other palynomorphs from the western and eastern margins
of the Labrador–Baffin Seaway.
Geological Survey of Denmark and Greenland Bulletin 36, 143 pp.
Bulletin36.qxp_Bulletin 36 19/12/16 13.39 Side 13
14
r
CanadaCanada
Ellesmere
Island
70°N
60°N
45°W55°W65°W
Labrado
r
Nuussuaq Nuussuaq
Basin
Hareøen
Disko
Bylot
Island
Cape
Dyer
Scott Inlet
Buchan Gulf
Quqaluit/
Padloping
islands
Saglek B
asin
646
112
113
645
Umiivik-1
Delta-1
T4-1
T8-1
Alpha-1/S1
Gamma-1
Hellefisk-1
AT7-1
AT2-1
LF7-1
Ikermiut-1
Nukik-2
Nukik-1
Qulleq-1
Gjoa G-37
Ralegh N-18
Hekja O-71
Rut H-11
Gilbert F-53
Karlsefni A-13
Skolp E-07
Pothurst P-19
Ogmund E-72
Snorri J-90
Bjarni O-82
Bjarni H-81
Herjolf M-92
North Bjarni F-06
Roberval K-92
North Leif I-05
Cartier D-70
Hare Bay E-21
Freydis B-87
South Labrador N-79
Kangâmiut-1
647
HopedaleBasin
Cum
berland
Sound
H
om
e Bay
B
af f in
Is lan
d
Canada
D
a
v is S
t ra
i t
La
b
ra
d
o
r S
e
a
GRO#3
250 km
Baffin
Bay
Greenland
H
udson
Stra i t
75°W
Shallow cored borehole
Greenland–Canada border
ODP or DSDP borehole
Exploration well with gas shows
Exploration well
Exploration well with oil and
gas shows
500 km
Greenland
Labrador
Sea
Baffin
Bay
D
avis Strait
Nar
es
St
ra
it
B
a
ffin
Isla
n
d
60°N
60°W 40°W
70°N
80°N
Bulletin36.qxp_Bulletin 36 19/12/16 13.39 Side 14
15
Canada and Greenland are separated, from south to
north, by the Labrador Sea, the Davis Strait, Baffin Bay
(which we call collectively the Labrador–Baffin Seaway),
and the narrow Nares Strait (Fig. 1). The Labrador–
Baffin Seaway stretches from about 52°N on to 75°N, a
distance of roughly 2500 km. Timing for the geological
evolution of the seaway’s margin is based primarily on
biostratigraphic analyses from exploration wells drilled
between 1971 and 2000 on the Labrador Margin, off
West Greenland and in the Davis Strait, and from some
shallow cored boreholes drilled in Baffin Bay in the
1980s (Figs 1, 2). These drilling activities revealed thick
succes sions of Mesozoic–Cenozoic sediments on both
sides of the seaway, and the lithostratigraphy of these
has developed in tandem with the biostratigraphy.
Additional in formation comes from numerous Creta ce -
ous–Cenozoic outcrop sections in the Nuussuaq Basin
in West Green land, a few onshore sections in Labrador
and on Baffin and Bylot islands (Figs 1, 2), and from
ODP Leg 105, hole 645 in Baffin Bay (Fig. 1).
Although previous work on these materials has provid-
ed a good stratigraphic foundation for the Cretaceous–
Cenozoic of the region (Fig. 3), renewed petroleum
exploration interest in recent years has revealed gaps in
our knowledge and a need to correlate the western and
eastern margins of the seaway. Consequently, for the
past decade, the Geo logical Survey of Denmark and
Greenland (GEUS) and the Geological Survey of Cana -
da (Atlantic) (GSCA) have undertaken an exhaustive
paly nological study, based on palynological analysis of
more than 2000 well samples. The objectives of this
work have been to refine age control and palaeoenvi-
ronmental interpretations on a regional scale, with the
ultimate goal of contribu ting fundamentally to an
under standing of the geological history of the seaway
and its petroleum systems. A significant finding has been
that the orderly story of five regional unconformities
(Fig. 3) is not as clear-cut as has been previously thought
(see Fig. 4 and Nøhr-Hansen et al. 2016 for details).
Dinoflagellate cysts (dinocysts) are the primary paly-
nomorph group evaluated, but spores and pollen (mio -
spores), a fern microspore massula (Azolla), algal micro-
fossils and acritarchs have also been used. Our new age
determinations (Figs 5, 6, in pocket; see Nøhr-Hansen
et al. 2016 for further details) are more precise than
those of previous studies because of advances made in
refining stratigraphic ranges, especially of dinocysts
(e.g. Williams et al. 2004). Moreover, the number of
species for which we have detailed stratigraphic infor-
mation has increased immeasurably since early studies
in the region in the 1970s.
This paper focuses on the systematic treatment of
palynomorphs observed in the study, thereby under-
pinning the results presented in the companion bulletin
(Nøhr-Hansen et al. 2016), but also facilitating future
work in the region and further afield.
Introduction
Facing page:
Fig. 1. Map of the Labrador–Baffin Seaway showing the location
of relevant wells, boreholes and onshore localities; wells and bore-
holes shown in red were used in this study. The cored borehole
GGU 400712 is 2 km ENE of Umiivik-1; Annertuneq is on the
north coast of Nuussuaq. Inset map shows the regional context of
the Labrador–Baffin Seaway between Canada and Greenland.
Bulletin36.qxp_Bulletin 36 19/12/16 13.39 Side 15
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645
Rut H-11
Gilbert F-53 Karlsefni A-13
Pothurst P-19
Skolp E-07
Ogmund E-72
Snorri J-90
Bjarni O-82
Roberval K-92
North Leif I-05
Freydis B-87
Cartier D-70
Herjolf M-92
North Bjarni F-06
250 km
South Labrador N-79
647
Saglek
Basin
Hellefisk-1
Ikermiut-1
Nukik-2
Delta-1
T4-1
T8-1
Gamma-1
Alpha-1/S1
Nukik-1
Qulleq-1
LF7-1
AT2-1
AT7-1
Gjoa F-37
Ralegh N-18
Hekja O-71
Kangâmiut-1
Nuussuaq
Svartenhuk
Halvø
Disko
Cape
Dyer
Umiivik-1
GRO#3
45°W 35°W
Sediments, large basins (400–0 Ma)
Oceanic crust, Eocene
Oceanic crust, age uncertain
Basalts and intrusives (60–30 Ma)
Basement locally covered
by sediment
Spreading axis
Extensional fault
Compressional fault
thrust
Transition fault
ODP or DSDP borehole
Oceanic crust, Paleocene
Exploration well
Exploration well with gas shows,
Exploration well with oil and
gas shows
Shallow cored borehole
Fault (undifferentiated)
Paam
iut B
asin
N
u
u
k
B
as
in
K
an
gâ
m
iu
t
B
as
in
Lady Franklin Basin
Sisimiut
Basin
Disko
Bugt
Ik
er
m
iu
t
B
as
in
H
o
m
e B
ay
B
asin
Lancaster
Basin
Eclipse
Trough
M
elville Bay
G
raben
Buchan
G
raben
Scott
G
raben
D
av
is
S
tr
ai
t
H
ig
h
Baffin
Basin
N
u
u
ss
u
au
q
B
as
in
Labrado
r
Labrador
Sea
Hudson Strait
C
um
berland Sound
M
elville Bay
K
ivioq
Basin
Canada
Greenland
Bylot
Island
Baffin
Island
65°W 55°W 45°W
60°N
70°
70°N
55°W65°W75°W
H
opedale
Basin
Hawke
Basin
West Greenland
Volcanic Province
Bulletin36.qxp_Bulletin 36 19/12/16 13.39 Side 16
17
The study is based on samples from ditch cuttings,
sidewall cores and conventional cores from onshore and
offshore wells and boreholes, and on outcrop samples
(Figs 1, 2). Most of the samples were processed for paly-
nology between 1990 and 2006, using standard techniques
for concentrating palynomorphs. However, many of the
sidewall-core samples are represented only by oil-company
slides produced in the late 1970s to 1980s and are of
variable quality.
Using transmitted light microscopy, qualitative and
quantitative analyses were carried out of the dinocysts,
acritarchs and pollen and spores, the results and impli-
cations of which are detailed in Nøhr-Hansen et al.
(2016). The systematic section below is accompanied
by plates; sample number, slide number and England
Finder coordinates for each specimen are provided. The
slides from the six offshore West Greenland wells,
Hellefisk-1, Ikermiut-1, Kangâmiut-1, Nukik-1, Nukik-
2 and Qulleq-1 are kept at GEUS. GEUS also processed
samples from six offshore Canadian wells, Gjoa G-37,
Hekja O-71, North Leif I-05, Ogmund E-72, Ralegh
N-18 and Skolp E-07. One set of palynology slides from
Gjoa G-37, Hekja O-71 and Ralegh N-18 is housed at
GEUS (Copenhagen) and one set is housed at the
Geological Survey of Canada (Atlantic), Dartmouth,
Nova Scotia, Canada. A set from each well is also curated
at the Canada–Nova Scotia Offshore Petroleum Board,
Dartmouth, Nova Scotia. The Geological Survey of
Canada has processed samples from all the Labrador
Margin wells, which include Bjarni O-82, Gilbert F-53,
Kalsefni A-13, Roberval K-92, Rut H-11, Snorri J-90
and South Labrador N-79. One set of palynology slides
from these wells is housed at the Geological Survey of
Canada (Atlantic), Dartmouth, Nova Scotia. Another
set is curated at the Canada – Newfoundland and
Labrador Offshore Petroleum Board (CNLOPB) in St.
Johns, Newfoundland. Also on file with CNLOPB are
sets of GEUS-prepared slides from North Leif I-05,
Ogmund E-72 and Skolp E-07.
Facing page:
Fig. 2. Map of the basic geology of the Labrador–Baffin Seaway,
showing the main structural features and relevant well/borehole
locations; wells and boreholes shown in red were used in this study.
Modified from Henriksen et al. (2009) and Oakey & Chalmers
(2012).
Methodology
Bulletin36.qxp_Bulletin 36 19/12/16 13.39 Side 17
18
Upper
Cretaceous
Paleocene
Eocene
Oligocene
Miocene
Pliocene
Quaternary
Lower
Cretaceous
0
Ma
5
10
15
20
25
30
35
40
45
50
55
60
65
70
75
80
85
90
95
100
105
110
115
120
125
130
135
140
145
Aptian
Barremian
Hauterivian
Valanginian
Berriasian
Albian
Cenomanian
Turonian
Coniacian
Santonian
Campanian
Maastrichtian
Danian
Selandian
Thanetian
Ypresian
Lutetian
Bartonian
Priabonian
Rupelian
Chattian
Aquitanian
Burdigalian
Langhian
Serravallian
Tortonian
Messinian
Zanclean
Piacenzian
C28
C29
C30
C31
C32
C33
C34n
C34n
C34n
C34n
M"-3"r
M"-2"r
M"-1"r
C27
C26
C25
C24
C22
C21
C20
C19
C18
C17
C23
C16
C15
C13
Magnetostrat.
(Polarity, Chron)
Chronostratigraphy Uncon-
formitiesShelf Basin
Labrador MarginSW NE
Labrador
Trough erosion
Leif
Mb
upper
Freydis Mb
lower
Freydis Mb
upper and
middle
Gudrid mb
lower Gudrid mb
Sa
gle
k F
m
Mokami Fm
lower
Kenamu Fm
Cartwright Fm
lower
lower
upper
upper
upper
Bjarni Fm
Alexis Fm
Markland Fm
(Labrador U
nc.)
(Avalon Unc.)
(Bylot Unc.)
Bylot
Avalon
Labrador
Baffin Bay
Beaufort
(Baffin Bay Unc.)
(B
eaufort U
n
c.
)
Fig. 3. Stratigraphic framework of
the Mesozoic–Cenozoic rocks of
the Labrador and West Green -
land margins and adjacent on -
shore sections; modified from
Gregersen et al. (2013). The La -
brador Margin stratigraphy is
from Dickie et al. (2011), with
unconformities from McWhae
(1981). The West Greenland
Mar gin stratigraphy is based on
well stratigraphic studies by Rolle
(1985), Nøhr-Hansen (2003) and
Sønderholm et al. (2003); the
deeper sub-well section is based
on Chalmers et al. (1993), Chal -
mers & Pulvertaft (2001) and
Sørensen (2006). The Nuussuaq
Basin stratigraphy is from Storey
et al. (1998), Dam et al. (2009),
Pedersen & Nøhr-Hansen (2014)
and Larsen et al. (2015); these pa -
pers and this study are the source
of the inferred ages. The south-
east Baffin Island stratigraphy is
from Burden & Langille (1990)
and Pedersen et al. (2002), the
Home Bay and Scott Inlet seabed
samples from MacLean et al.
(2014) and the north Baffin Is -
land stratigraphy is based on Jack -
son et al. (1978) and McWhae
(1981). The Bylot Island strati -
graphy is from Miall (1986),
Waterfield (1989), and Harrison
et al. (1999). The timescale (Ma)
and magnetostratigraphy are from
Gradstein et al. (2012). Fm: For -
mation (formal). fm: formation
(informal). Mb: Member (for -
mal). mb: member (informal).
Unc.: unconformity.
Bulletin36.qxp_Bulletin 36 19/12/16 13.39 Side 18
19
H
el
le
fis
k-
1
G
R
O
#
3
U
m
iiv
ik
-1
Ik
er
m
iu
t-
1
K
an
gâ
m
iu
t-
1
N
u
ki
k-
2
N
u
ki
k-
1
Q
u
lle
q
-1
West Greenland Margin
Kangilia Fm
Nuussuaq Basin SE Baffin IslandNWSES N Bylot Island
Itilli Fm
Aaffarsuaq
Mb
Atane Fm
?
(Ikermiut Fm)
Kangeq sequence
Ikermiut Fm
Ikermiut Fm
Narssarmiut Fm
Kangâmiut Fm
Kangâmiut Fm
Kangâmiut Fm
Kangâmiut
Fm
Hellefisk FmNukik Fm
Nukik Fm
?
?
?
?
?
? ?
? ?
? ?
Ataneq
Fm
Ataneq Fm
Manîtsoq Fm
Manîtsoq Fm
Manîtsoq Fm
Fylla sand
Erqua Fm
Naqerlog Fm
Hareøen Fm
(Hareøen)
Youngest
volcanics
Quqaluit Fm
Cape Searle Fm
N
u
u
ss
u
aq
G
ro
u
p
(Home Bay –
Scott Inlet)
Cape Dyer Basalt
Maligât Fm
Svartenhuk Fm
Vaigat Fm
Atanikerluk Fm
Eqalulik Fm
Quikavsak Fm
Kome Fm
Basalts?
Sills/dykes
Appat sequence
Kitsissut
sequence
?
?
Upernivik
Næs Fm
Slibestens-
fjeldet Fm
Hassel Fm
(Labrador Unc.)
Sermilik fm
Kanguk Fm
Aktineq fm
Pond Inlet fm
Navy Board fm
Bylot Island fm
Agatdal Fm
Ikermiut Fm
Mainly marine
mudstones, locally
sandy/silty
Mainly marine or
deltaic sandy/silty
deposits, locally
mudstones
Potential and
known source rocks
Volcanics
TD in basement
Unconformity
Mainly continental
deposits
Bulletin36.qxp_Bulletin 36 19/12/16 13.39 Side 19
20
0
Ma
5
10
15
20
25
30
35
40
45
50
55
60
65
70
75
80
85
90
95
100
105
110
115
120
125
Late
Cretaceous
Paleocene
Eocene
Oligocene
Miocene
Pliocene
Quaternary
P
er
io
d
/
E
p
o
ch
A
ge
(
st
ag
e)
Fo
rm
at
io
n
M
em
b
er
R
eg
io
n
al
(D
ic
ki
e
et
a
l.
2
0
1
1
)
L
ab
ra
d
o
r
M
ar
gi
n
(M
cW
h
ae
1
9
8
1
)
Early
Cretaceous
Aptian
Albian
Cenomanian
Turonian
Coniacian
Santonian
Campanian
Maastrichtian
Danian
Selandian
Thanetian
Ypresian
Lutetian
Bartonian
Priabonian
Rupelian
Chattian
Aquitanian
Burdigalian
Langhian
Serravallian
Tortonian
Messinian
Zanclean
Piacenzian
Mokami
Kenamu
Cartwright
Markland
Bjarni
Saglek
Upper
Lower
Upper
Leif
Lower
Lower
Upper +
Middle Gudrid
Lower
Gudrid
Upper
Freydis
Lower
Freydis
Upper
Sn
o
rr
i
J-
9
0
H
e
k
ja
O
-7
1
N
o
rt
h
L
e
if
I-
0
5
B
e
au
fo
rt
B
af
fin
B
ay
B
yl
o
t
A
va
lo
n
L
ab
ra
d
o
r
R
o
b
e
rv
al
K
-9
2
B
ja
rn
i
O
-8
2
O
gm
u
n
d
E
-7
2
S.
L
ab
ra
d
o
r
N
-7
9
Sk
o
lp
E
-0
7
K
ar
ls
e
fn
i
A
-1
3
G
ilb
e
rt
F
-5
3
R
u
t
H
-1
1
R
al
e
gh
N
-1
8
G
jo
a
G
-3
7
S N
Chronostratigraphy Hiatuses
Labrador Margin
Preserved stratigraphic interval
Labrador Margin
lithostratigraphy
Bulletin36.qxp_Bulletin 36 19/12/16 13.39 Side 20
21
Ma
H
e
lle
fis
k
-1
N
u
k
ik
-1
N
u
k
ik
-2
K
an
gâ
m
iu
t-
1
Ik
e
rm
iu
t-
1
Q
u
lle
q
-1
G
R
O
#
3
&
s
u
rf
ac
e
se
ct
io
n
s
N
.
N
u
u
ss
u
aq
U
m
iiv
ik
-1
,
Sv
ar
te
n
h
u
k
H
al
vø
&
N
.
N
u
u
ss
u
aq
,
su
rf
ac
e
s
e
ct
io
n
s
S N
Late
Cretaceous
Paleocene
Eocene
Oligocene
Miocene
Pliocene
Quaternary
P
er
io
d
/
E
p
o
ch
A
ge
(
st
ag
e)
Early
Cretaceous
Aptian
Albian
Cenomanian
Turonian
Coniacian
Santonian
Campanian
Maastrichtian
Danian
Selandian
Thanetian
Ypresian
Lutetian
Bartonian
Priabonian
Rupelian
Chattian
Aquitanian
Burdigalian
Langhian
Serravallian
Tortonian
Messinian
Zanclean
0
5
10
15
20
25
30
35
40
45
50
55
60
65
70
75
80
85
90
95
100
105
110
115
120
125
N
o
t
p
en
et
ra
te
d
o
ff
sh
o
re
Chronostratigraphy
West Greenland Margin
Preserved stratigraphic interval
Stratigraphic
record
Labrador
0
% %
100 100
West
Greenland
(offshore)
0
Piacenzian
Fig. 4. Ages of strata preserved in
wells on the Labrador, Baffin and
Green land Margins, compared
with the hiatuses linked to
uncon formities reported in the
literature. Dickie et al. (2011)
recognised a number of uncon-
formities in two or more of three
regions – the Southwest Greenland
Shelf, the Labrador Shelf and the
Jeanne d’Arc Basin, offshore
Newfound land (red, all three
regions; green, two regions).
McWhae (1981) named five
unconformities on the Labrador
Margin; the hiatuses associated
with these unconformities indi-
cated here are estimates by
McWhae (1981) based on an
unspecified timescale and thus
should be considered approxi-
mate. The ‘cumulative preserva-
tion’ plots (right) depict sche ma-
tically the proportion of wells
that contain sediments of a speci-
fied age, based on one million-
year slots; note that these cal -
culations only include wells that
extend down to/beyond the rele-
vant stratigraphic level.
Bulletin36.qxp_Bulletin 36 19/12/16 13.39 Side 21
22
Phelodinium
Apectodinium,Axiodinium, Charlesdowniea,
Petalodinium, Piladinium, Rhombodinium,
Sophismatia, Stichodinium, Talladinium, Wetzeliella
Alterbidinium, Cerodinium, Chatangiella, Deflandrea,
Isabelidinium, Lentinia, Palaeocystodinium,
Scalenodinium?, Senegalinium, Spinidinium,
Trithyrodinium
Ginginodinium?, Laciniadinium,
Palaeoperidinium, Phthanoperidinium?
Subtilisphaera
Atopodinium, Batiacasphaera, Chlamydophorella,
Impletosphaeridium?, Raphidodinium,
Reticulatosphaera, Simplicidinium,
Tanyosphaeridium, Wallodinium
Alisocysta, Eocladopyxis, Homotryblium
Hystrichosphaeridium
Heteraulacacysta
Tuberculodinium
Aptea, Nyktericysta, Odontochitina, Pseudoceratium,
Taurodinium, Vesperopsis, Xenascus
Adnatosphaeridium, Areoligera, Areosphaeridium,
Chiropteridium, Cleistosphaeridium, Enneadocysta,
Glaphyrocysta, Heterosphaeridium, Licracysta
Palynodinium, Schematophora, Senonaisphaera,
Tenua
Callaiosphaeridium, Cerebrocysta,
Chytroeisphaeridia, Fibrocysta, Habibacysta,
Kiokansium, Surculosphaeridium, Trichodinium
Cannosphaeropsis, Evittosphaerula,
Hystrichosphaeropsis, Hystrichostrogylon,
Impagidinium, Rottnestia, Spiniferites,
Achilleodinium, Apteodinium, Cordosphaeridium,
Cribroperidinium, Dapsilidinium, Diphyes,
Disphaerogena, Hapsocysta, Hystrichokolpoma,
Lingulodinium, Operculodinium, Spongodinium,
Thalassiphora
Eatonicysta, Kleithriasphaeridium,
Oligosphaeridium
Gillinia, Histiocysta
Batioladinium
Uncertain
Cribroperidinioideae
Gonyaulacoideae
Uncertain
Helgolandinioideae
Goniodomoideae
Protoperidinioideae
Podolampaceae
Peridiniaceae
Ceratiaceae
Ceratocoryaceae
Areoligeraceae
Heterocapsaceae
Heterodiniaceae
Uncertain
Crypthecodiniaceae
Pyrocystaceae
Goniodomaceae
Pyrodinioideae
Gambierdiscoideae
Glenodiniaceae
Palaeoperidinioideae
Deflandreoideae
Calciodinelloideae
Ovoidinioideae
Lithoperidinioideae
Peridinioideae
Wetzelielloideae
Leptodinioideae
Diplopsalioideae
Protoperidinioideae
Gonyaulacaceae
Rhaetogonyaulacaceae
Cladopyxiaceae
Pareodiniaceae
Pareodinioideae
Broomeoideae
O
rd
er
G
o
n
ya
u
la
ca
le
s
O
rd
er
P
er
id
in
al
es
Peridiniineae
Ceratiineae
Gonyaulacineae
Uncertain
Goniodomineae
Heterocapsineae
Glenodiniineae
Cladopyxiineae
Rhaetogonyaulacineae
Family Subfamily GeneraSuborder
Fig. 7. Suprageneric affiliation of peridiniphycidean genera treated systematically in the present study. The only non-peridiniphycidean dinocyst
genus treated in this study is Dinogymnium, which belongs to the subclass Gymnodiniophycidae, order Ptychodiscales, family Ptychodiscaceae,
subfamily Dinogymnioideae.
Bulletin36.qxp_Bulletin 36 19/12/16 13.39 Side 22
23
Division Dinoflagellata (Bütschli 1885)
Fensome et al. 1993
Class Dinophyceae Pascher 1914
Genus Achilleodinium Eaton 1976
Type. Eisenack 1954, plate 11, fig. 18, as Hystricho -
sphaer idium biformoides.
1976 Achilleodinium Eaton, p. 234.
Remarks. We follow the synopsis of Fensome et al.
(2009, p. 11) for this genus.
Achilleodinium biformoides (Eisenack 1954)
Eaton 1976
(Plate 1, figs 1–3)
1954 Hystrichosphaeridium biformoides Eisenack,
p. 68, plate 11, figs 16–20.
1965 Baltisphaeridium biformoides (Eisenack) –
Downie & Sarjeant, p. 87.
1965 Hystrichokolpoma biformoides (Eisenack) –
Rozen, p. 308.
1976 Achilleodinium biformoides (Eisenack) –
Eaton, p. 234.
1980 Florentinia biformoides (Eisenack) – Duxbury,
p. 121.
Age. LO: Ypresian.
Genus Adnatosphaeridium Williams & Downie
1966a
Type. Williams & Downie 1966a, plate 24, fig. 7, text-
fig. 56, as Adnatosphaeridium vittatum.
1966a Adnatosphaeridium Williams & Downie,
p. 215.
Remarks. Stancliffe & Sarjeant (1990, p. 199–200) emen -
ded the diagnosis of Adnatosphaeridium but did not change
the concept of the generic circumscription. Here, we follow
the synopsis provided by Fensome et al. (2009, p. 11).
Systematics – general
Systematics – dinoflagellates
Presentation of the systematic taxonomic descriptions
includes subsections on dinocysts, acritarchs and other
algae, miospores, and fungi. For pragmatic reasons,
genera are arranged alphabetically within each section.
For the dinocysts, a tabulation of suprageneric affinities
is provided (Fig. 7). Under individual taxa, all synonyms
are listed but those already indicated and credited to
earlier works in Fensome et al. (2008) are generally not
discussed further. The term ‘synopsis’ is used for a state-
ment of what we consider the essential defining features
of the genus. Unless otherwise clearly indicated, the
synopses given herein should not be understood to repre-
sent emendations. Only those emendations that we agree
with or consider helpful are cited.
All ages refer to their definition in the latest revision of
the geological timescale (Gradstein et al. 2012). For most
species, an age is indicated for significant events in its
stratigraphical range, usually a range top or ‘Last
Occurrence’ (LO). The statement ‘not plotted’ refers to
the fact that a particular taxon was not plotted in the
summary event charts (Figs 5, 6, in pocket); the reasons
for not plotting some taxa include a taxon’s sparsity or
long and unhelpful range (though it may be helpful
palaeoenvironmentally. Because the data are based main -
ly on cuttings samples, a taxon’s range bottom or ‘First
Occurrence’ (FO) is rarely used. We occasionally refer to
the LAD of a species, its ‘Last Appearance Datum’,
which refers to its youngest global occurrence.
Bulletin36.qxp_Bulletin 36 19/12/16 13.39 Side 23
24
Adnatosphaeridium vittatum Williams & Downie
1966a
(Plate 1, figs 4, 8)
1966a Adnatosphaeridium vittatum Williams &
Downie, p. 15, plate 24, fig. 7; text-fig. 56.
1966a Adnatosphaeridium multispinosum Williams &
Downie, p. 216, 217, plate 24, fig. 5; text-fig.
57.
Age. LO: late Bartonian. Not plotted.
Remarks. Following Fensome et al. (2009, p. 13), the
species Adnatosphaeridium multispinosum is considered
a taxonomic synonym of Adnatosphaeridium vittatum.
Genus Alisocysta Stover & Evitt 1978
Type. Drugg 1967, plate 1, fig. 12, as Eisenackia cir -
cum tabulata.
1978 Alisocysta Stover & Evitt, p. 15, 16.
1979a Agerasphaera Harland, p. 28, 29; illegitimate
name, having the same type as Alisocysta.
Synopsis. Goniodomacean (pyrodinioid) cysts that are
proximate and subspherical. The tabulation is reflected
by penitabular ridges or septa that delineate the plates,
including the cingulars and some or all of the sulcals.
The archaeopyle is apical, with the formula A(1–4´);
operculum free.
Remarks. Quattrocchio & Sarjeant (2003, p. 144) consid-
ered Alisocysta to be a taxonomic junior synonym of
Eisenackia. However, as stated by Stover & Evitt (1978,
p. 42) “Alisocysta has parasutural [i.e. penitabular] ridges
or septa rather than depressions as in Eisenackia” and
Alisocysta is thus retained here as a separate genus, includ-
ing both the type, Alisocysta circumtabulata, and Alisocysta
margarita.
Alisocysta circumtabulata (Drugg 1967) Stover &
Evitt 1978
(Plate 1, figs 6, 7)
1967 Eisenackia circumtabulata Drugg, p. 15, plate
1, figs 12, 13.
1978 Hystrichokolpoma circumtabulatum (Drugg) –
Schumacker-Lambry, p. 42.
1978 Alisocysta circumtabulata (Drugg) – Stover &
Evitt, p. 16.
1979a Agerasphaera circumtabulata (Drugg) –
Harland, p. 29; illegitimate combination as
the generic name Agerasphaera is illegitimate.
2003 Eisenackia circumtabulata (Drugg) –
Quattrocchio & Sarjeant, p. 146.
Age. LO: earliest Thanetian.
Alisocysta margarita (Harland 1979a) Harland
1979a
(Plate 1, figs 5, 9)
1979a Agerasphaera margarita Harland, p. 29, 31,
33, plate 1, figs 1–12; plate 2, figs 1–10.
1979a Alisocysta margarita (Harland) – Harland,
p. 35.
2003 Eisenackia margarita (Harland) –
Quattrocchio & Sarjeant, p. 146.
Age. LO: earliest Thanetian.
Genus Alterbidinium Lentin & Williams 1985
emend. nov.
Type. Vozzhennikova 1967, plate 77, fig. 2, as Albertia
recticornis.
1967 Albertia Vozzhennikova, p. 150, 151; illegiti-
mate name.
1976 Alterbia Lentin & Williams, p. 47, 48; illegiti-
mate name.
1985 Alterbidinium Lentin & Williams, p. 14.
Emended description. Peridiniacean (deflandreoid) cysts
that are proximate and peridinioid, usually elongate, in
outline. The antapical horns are always asymmetrically
arranged, the left horn being larger. Bicavate. The peri-
cyst surface is generally atabulate, smooth, or with low
ornament; the cingulum is commonly indicated, if only
marginally. The periarchaeopyle is intercalary or combi-
nation intercalary–precingular; always involving an iso-
to stenodeltaform hexa plate 2a and commonly plate
4´´, the operculum remaining attached posteriorly;
archaeopyle I2a @ or (I2aP4´´)@.
Remarks. In their emendation of Alterbidinium, Khowaja-
Ateequzzaman et al. (1991, p. 38) stated: “archaeopyle
intercalary, independently developed on periphragm and
endophragm, dissimilar in shape; periarchaeopyle hexa
2a, steno/iso-deltaform, perioperculum free or adnate;
endoarchaeopyle hexa 2a, eury-deltaform, endoperculum
adnate.” However, they did not base their emendation on
Bulletin36.qxp_Bulletin 36 19/12/16 13.39 Side 24
the type of the genus. Thus, in the emendation proposed
here, the synop sis of Fensome et al. (2009, p. 13) is largely
repeated: it emphasised the nature of the archaeopyle,
which is formed from the iso- to stenodeltaform hexa
plate 2a, but may also involve plate 4´´. Almost invari-
ably, the operculum remains attached posteriorly.
Genera with similar morphologies to Alterbidinium
include Spinidinium, Diconodinium, Cerodinium, Cha -
tan giella and Isabelidinium. Spinidinium and Dicono -
dinium differ from Alterbidinium in having a spinate
peri ph ragm. Cerodinium has a large isodeltaform periar-
chaeopyle, an endoarchaeopyle formed from the loss of one
to three anterior intercalary plates, and generally long apical
and antapical horns. Chatangiella is distinguished by its
gen erally omegaform 2a and partite cingulum, although
Fen some et al. (2009, p. 19) stated that the archaeopyle
could vary between isodeltaform and iso-omegaform. We
now consider that Chatan giella should be restricted to taxa
that possess an iso- or lati-omegaform archaeopyle. Sim -
ilarly, Isabelidinium is restricted to taxa with an iso- to lati-
omega form 2a plate and archaeopyle. However, some
species presently included in Isabelidinium can have the -
taform or isodeltaform 2a plates and archaeopyles.
The present morphological differences used to separate
the deflandreoid genera are in large part unsatisfactory.
Perhaps a more effective approach would be to adopt the
methodological approach used by Williams et al. (2015) in
their reclassification of the wetzelielloideans. Following this
example, the deflandroidean genera would be differentiated
primarily on the nature of the archaeopyle, since this
reflects differences in tabulation pattern. This then follows
the categories recognised by Lentin & Williams (1976) and
expanded by Bujak & Davies (1983). These are omega -
form, thetaform and deltaform modes, with steno-, iso-
and lati- subcategories. Such an approach would not be a
perfect solution as there are clearly gradations, but it would
make the generic distinctions more meaningful, and is
considered here the best solution.
In the spirit of this approach, herein we informally use
Isabelidinium for forms having a lati- to iso-omegaform
archaeopyle. A formal revision would require many of the
species now included in that genus to be transferred to
other existing or new genera. Indeed, a re-appraisal of all
deflandreoidean taxa is deemed necessary, but is beyond the
scope of the present paper, although as an initial step emen-
dations are proposed for Alterbidinium and Chatangiella.
Alterbidinium acutulum (Wilson 1967a) Lentin &
Williams 1985
(Plate 1, figs 10, 11)
1967a Deflandrea acutula Wilson, p. 225, 226, figs
11, 12.
1967 Albertia curvicornis Vozzhennikova, p. 151,
plate 76, figs 1–4.
1967 Albertia recticornis Vozzhennikova, p. 151,
152, plate 77, figs 1–4; plate 78, figs 1–3;
plate 79, figs 1, 2.
1976 Alterbia acutula (Wilson) – Lentin &
Williams, p. 48.
1976 Alterbia curvicornis (Vozzhennikova) – Lentin
& Williams, p. 49.
1976 Alterbia recticornis (Vozzhennikova) – Lentin
& Williams, p. 47.
1985 Alterbidinium acutulum (Wilson) – Lentin &
Williams, p. 14.
Age. LO: early Maastrichtian.
Remarks. Fensome & Williams (2005, p. 9) recorded this
species, in part, under the informal name ‘Alterbidinium
fleximorphum’ because of the great variation shown by
this species. It is now recognised that this variability
should be encompassed within Alterbidinium acutulum.
Alterbidinium biaperturum (McIntyre 1975) comb.
nov.
(Plate 1, figs 12–14)
1975 Deflandrea biapertura McIntyre, p. 66, plate
3, figs 5–8.
1976 Chatangiella? biapertura (McIntyre) – Lentin
& Williams, p. 53.
Age. LO: late Maastrichtian.
Remarks. This species is distinguished by an opening in
the pericyst between the antapical horns. From the illus-
trations of the holotype in McIntyre (1975, plate 3, figs
5, 6), Alterbidinium (as Chatangiella?) biaperturum ap -
pears to possess a cingulum delineated by continuous
ridges; the periarchaeopyle is extremely unusual, being
distinctly deltaform on one lateral margin but, as M.
Pearce (personal communication 2015) has noted, possi-
bly thetaform on the other lateral margin. Whether such
asymmetry occurs in other specimens of this species is
unknown. But regardless, the species is thus not assign -
able to Chatangiella. Lentin & Williams (1976) were
aware of these differences when provisionally including
this taxon in Chatangiella, but did not consider assigning
it to Alterbidinium. We herein transfer the species to
Alter bidinium to more accurately reflect its morphology.
25
Bulletin36.qxp_Bulletin 36 19/12/16 13.39 Side 25
26
Alterbidinium? bicellulum (Islam 1983a) Lentin &
Williams 1985
(Plate 1, figs 15, 16)
1983a Alterbia? bicellula Islam, p. 335, 336, plate 1,
figs 6, 7.
1985 Alterbidinium? bicellulum (Islam) – Lentin &
Williams, p. 14.
Age. LO: Lutetian, with a peak occurrence in the
Ypresian.
Remarks. In the diagnosis for Alterbidinium? bicellulum
(as Alterbia? bicellula), Islam (1983a, p. 336) stated that
the archaeopyle was “… intercalary type I/I with stan-
dard hexa style operculum attached”. This implies that
the archaeopyle is isodeltaform, but it is not possible to
confirm this from his illustrations (Islam 1983a, plate 1,
figs 6, 7, 10, 11). Islam (1983a) questionably assigned
this species to Alterbia (now Alterbidinium) “because the
epipericoel is not always communicative to the exterior,
and its degree of cavation and the archaeopyle index do
not match those prescribed for the genus. The degree of
cavation and the archaeopyle index of this species also do
not match those of other peridinioid genera that are
differentiated on the basis of these features by Lentin &
Williams (1976).” Taxa with an isodeltaform archaeo -
pyle and an attached operculum are rare, especially in the
Cenozoic. Thus we leave the species questionably in
Alberbidinium.
Alterbidinium ioannidesii Pearce 2010
(Plate 1, Fig. 17)
1986 Dinoflagellate type E of Ioannides, p. 42,
plate 23, figs 13–16.
2010 Alterbidinium ioannidesii Pearce, p. 66, 67,
plate 1, figs 1–6.
Age. LO: early Campanian.
Remarks: This is an unusual species of Alterbidinium in
displaying tabulation on the pericyst.
Alterbidinium varium Kirsch 1991
(Plate 1, figs 18, 19)
1991 Alterbidinium varium Kirsch, p. 98, 99, plate
19, figs 1–10; text-figs 46a–h, 47a, b.
Age. LO: Campanian.
Genus Apectodinium (Costa & Downie 1976)
Lentin & Williams 1977a emend. Williams,
Damassa, Fensome & Guerstein in Fensome et al.
2009
Type. Deflandre & Cookson 1955, plate 5, fig. 7, as
Wetzeliella homomorpha.
1976 Wetzeliella subgenus Apectodinium Costa &
Downie, p. 608.
1977a Apectodinium (Costa & Downie 1976) –
Lentin & Williams, p. 8.
2009 Apectodinium (Costa & Downie) – emend.
Williams, Damassa, Fensome & Guerstein in
Fensome et al., p. 13, 14.
Remarks. The emended diagnosis in Fensome et al. (2009,
p. 13, 14) is followed here; this emphasised archaeopyle
type and variability and the general absence of a pericoel
except in the vicinity of the horns.
Apectodinium homomorphum (Deflandre &
Cookson 1955) Lentin & Williams 1977a
(Plate 1, fig. 20)
1955 Wetzeliella homomorpha Deflandre &
Cookson, p. 254, plate 5, fig. 7; text-fig. 19.
1974 Hystrichosphaeridium caiobense Regali et al., p.
290, plate 24, fig. 4.
1977a Apectodinium homomorphum (Deflandre &
Cookson) – Lentin & Williams, p. 8.
1981 Apectodinium caiobense (Regali et al.) – Lentin
& Williams, p. 14.
1983a Apectodinium folliculum Islam, p. 336, 337,
plate 1, figs 8, 9.
Age. Peak: early Ypresian.
Remarks. Harland (1979b, p. 64) emended the diagnosis
of this species, as Wetzeliella (Apectodinium) homomor-
phum, to “take into account the nature of the archaeo -
pyle, cingular and sulcal details”. A characteristic feature
of Apectodinium homomorphum is the absence of horns,
although Harland noted that incipient horns may be
developed in the apical, lateral and antapical areas.
Apectodinium parvum (Alberti 1961) Lentin &
Williams 1977a
(Plate 2, figs 1–3)
1961 Wetzeliella parva Alberti, p. 8, 9, plate 1, figs
14–18; plate 12, figs 10–12.
Bulletin36.qxp_Bulletin 36 19/12/16 13.39 Side 26
27
1976 Wetzeliella subgenus Apectodinium parva
(Alberti) – Costa & Downie, p. 608.
1977a Apectodinium parvum (Alberti) – Lentin &
Williams, p. 9.
Age. LO: earliest Ypresian.
Remarks. In his emendation of Apectodinium parvum,
Harland (1979b, p. 65, 66) noted that lateral horns are
absent and the processes may be intratabular or sutural.
In his description he stated that “The horn develop-
ment, prominent at the antapex, is variously developed
with relation to the apical horn, which may or may not
be present.” The Labrador Margin specimens usually
have an apical horn, but show considerable variation in
length and width. Some of the Labrador Margin speci-
mens have longer processes than those illustrated by
Harland (1979b), with those of one specimen being 21
μm. When longer, the processes are extremely delicate.
However, forms with more robust processes also occur.
Apectodinium quinquelatum (Williams & Downie
1966b) Costa & Downie 1979
(Plate 2, fig. 4)
1948 Hystrichosphaeridium geometricum Pastiels, p.
41, plate 4, figs 1–5, 7–10, non Hystricho -
sphaeridium geometricum Deflandre 1942.
1966b Wetzeliella homomorphum var. quinquelata
Williams & Downie, p. 191, 192, plate 18,
fig. 7.
1977a Wetzeliella homomorpha subsp. quinquelata
(Williams & Downie) – Lentin & Williams,
p. 8.
1979b Wetzeliella (Apectodinium) quinquelata
(Williams & Downie) – Harland, p. 67.
1979 Apectodinium quinquelatum (Williams &
Downie) – Costa & Downie, p. 43.
Age. LO: Ypresian. Not plotted.
Genus Aptea Eisenack 1958
Type. Eisenack 1958, plate 22, fig. 5, as Aptea polymorpha.
1958 Aptea Eisenack, p. 393.
1966a Doidyx Sarjeant, p. 205.
Aptea polymorpha Eisenack 1958
(Plate 2, fig. 5)
1958 Aptea polymorpha Eisenack, p. 394, plate 22,
figs 5–12; plate 24, fig. 5.
1986 Pseudoceratium polymorphum (Eisenack 1958)
– Bint, p.145.
Age. LO: Aptian.
Remarks. The ornamentation of this species is variable,
ranging from coarsely reticulate to irregularly spinate.
Genus Apteodinium Eisenack 1958
Type. Eisenack 1958, plate 23, fig. 9, as Apteodinium
granulatum.
1958 Apteodinium Eisenack, p. 385.
1961 Emslandia Gerlach, p. 171.
1971 Coniferatium Burgess, p. 80, 81.
Remarks. The synopsis for Apteodinium followed here is
that provided in Fensome et al. (2009, p. 14), which
noted that tabulation may sometimes be weakly reflected
on the cyst wall.
Apteodinium australiense (Deflandre & Cookson
1955) Williams 1978
(Plate 2, fig. 6)
1955 Gymnodinium australiense Deflandre &
Cookson, p. 248, plate 5, fig. 1.
1961 Gonyaulax tenuitabulata Gerlach, p. 159,
plate 25, figs 10, 11; text-figs 1–3.
1965 Emslandia australiensis (Deflandre &
Cookson) – Nagy, p. 202; combination not
validly published.
1965a Scriniodinium australiense (Deflandre &
Cookson) – Cookson & Eisenack, p. 122.
1969 Gonyaulacysta tenuitabulata (Gerlach) – de
Coninck, p. 23.
1978 Apteodinium australiense (Deflandre &
Cookson) – Williams, p. 794.
1978 Millioudodinium tenuitabulatum (Gerlach) –
Stover & Evitt, p. 174.
1981 Emslandia crassimurata Benedek & Sarjeant,
p. 320, 322, fig. 1, nos 2, 4.
1984 Cribroperidinium tenuitabulatum (Gerlach) –
Helenes, p. 124.
1984a Rhynchodiniopsis tenuitabulata (Gerlach) –
Sarjeant, p. 76.
Age. LO: Rupelian.
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28
Remarks. Apteodinium australiense has a spongy wall, as
does the holotype of Cribroperidinium tenuitabulatum.
Moreover, the latter has at best weakly developed and
barely discernible parasutural ridges. Hence, we consider
Cribroperidinium tenuitabulatum to be a junior synonym
of Apteodinium australiense.
Apteodinium spiridoides Benedek 1972
(Plate 2, figs 7, 8)
1972 Apteodinium spiridoides Benedek, p. 5, plate 2,
fig. 1a, b; plate 15, figs 1–6.
1981 Emslandia spiridoides (Benedek) – Benedek &
Sarjeant, p. 318.
Age. LO: earliest Serravallian.
Remarks. In their emendation of Apteodinium spiridoides,
Benedek & Sarjeant (1981, p. 318, 319) noted the pres-
ence of an apical horn, the two-layered nature of the wall
and the archaeopyle type. Jan du Chêne et al. (1986, p.
48) retained the species in Apteodinium.
Genus Areoligera Lejeune-Carpentier 1938
Type. Lejeune-Carpentier 1938, text-fig. 2, as Areoli -
gera senonensis.
1938 Areoligera Lejeune-Carpentier, p. B164.
Remarks. We agree with the synopsis for Areoligera pro -
vided by Fensome et al. (2009, p. 14) and their observa-
tions under remarks. Included in Areoligera are several
species with overlapping morphology. This is primarily
manifested in the shape and distribution of the process
complexes and the complexity of the linkages between
processes within an individual complex. Fensome et al.
(2009, p. 15) introduced two terms for the distribution
of the process complexes. The encircling condition is
when complexes are, “…arcuate to annulate and occur
on all the pre- and postcingular plates.” In the disjunct
condition, “…annulate to arcuate complexes are restrict-
ed to the dorsal surface, there is lateral development of
linear complexes and the ventral surface (i.e. 6´´ and
6´´´) is devoid of complexes.” Fensome et al. (2009, p.
15) define Areoligera gippingensis as a species with “encir-
cling complex arrangement and predominantly trabecular
connections.” Following the specific differences re cognised
for species of Areoligera by these authors, it is concluded
that all specimens with this general morphology studied
here should be assigned to Areoligera gippingensis.
The genera Chiropteridium and Glaphyrocysta superfi-
cially resemble Areoligera, but neither Chiropteridium nor
Glapyrocysta have distinct annulate or arcuate process
complexes with basal ridges that delineate tabulation.
However, Chiropteridium gilbertii does have intratabular
processes or complexes. Chiropteridium differs from Areo -
ligera primarily in having neither mid-dorsal nor mid-
ventral processes or membranes. Glaphyrocysta has non -
tabular or indistinctly contabular processes.
Areoligera circumsenonensis Fensome et al. 2009
(Plate 2, fig. 9)
1966a Areoligera cf. senonensis of Williams &
Downie, p. 230, 231; text-fig. 64 A–C (not
plate 25, fig 6, typographically mislabelled
plate 26, fig. 6).
1969 Areoligera senonensis Lejeune-Carpentier –
Gocht, p. 56, plate 8, fig. 4a, b (?not figs 5–
9); text-figs 40a, b, ?40c, d (not fig. 40e, f).
2009 Areoligera circumsenonensis Fensome et al., p.
15, plate 1, fig. m.
Age. LO: early Lutetian. Not plotted.
Remarks. Areoligera circumsenonensis has processes like
those of Areoligera senonensis but the process complexes
are annular to arcuate on the large pre- and postcingular
plates. In Areoligera gippingensis, the processes are inter-
connected along their length within, and occasionally
between, complexes.
Areoligera gippingensis Jolley 1992
(Plate 2, figs 10, 11)
1966a Areoligera cf. medusettiformis of Williams &
Downie, p. 229, plate 25, fig. 4.
1976 Areoligera cf. medusettiformis of Eaton, p. 246,
plate 3, fig. 7.
1992 Areoligera gippingensis Jolley, p. 26, 28, 30,
31, plate 1, figs 1–6; plate 2, figs 1–6; text-
figs 2a–d, 3.
Age. LO: late Ypresian. Peak: early Thanetian.
Remarks. Specimens of Areoligera gippingensis in Labrador
Margin samples show considerable variability in the delin-
eation of the process complexes. We have found that this
species is gradational with Glaphyrocysta divaricata. The
species can be common in the late Thanetian, but gener-
ally peaks in the early Thanetian.
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29
Genus Areosphaeridium Eaton 1971
Type. Klumpp 1953, plate 18, figs 3, 4, as Hystricho -
sphaeridium dictyoplokum.
1971 Areosphaeridium Eaton, p. 357, 358.
Synopsis. Chorate gonyaulacinean cysts with subspheri-
cal central body. Acavate. Processes mesotabular, solid,
commonly fibroid, expanded clypeate and fenestrate to
reticulate distally. The number of cingular processes is
variable. Archaeopyle apical, with formula A(1 –4´), oper-
culum free.
Remarks. The synopsis is based on the emendation of
Areosphaeridium by Stover & Williams (1995, p. 100).
Following Fensome et al. (2007), Areosphaeridium is
considered an areoligeracean cyst related to Enneadocysta.
Areosphaeridium diktyoplokum (Klumpp 1953)
Eaton 1971
(Plate 2, fig. 12)
1953 Hystrichosphaeridium diktyoplokum Klumpp, p.
392, plate 18, figs 3–7 (not plate 18, figs 8–10,
which are now Cordosphaeridium latum).
1963a Cordosphaeridium diktyoplokum (Klumpp) –
Eisenack, p. 262.
1971 Areosphaeridium diktyoplokum (Klumpp) –
Eaton, p. 358, 359.
Age. LO: latest Priabonian.
Genus Atopodinium Drugg 1978
Type. Drugg 1978, plate 1, fig. 1, as Atopodinium pro -
statum.
1978 Atopodinium Drugg, p. 62.
1981 Maghrebinia Below, p. 22.
1987 Bejuia Stover & Williams, p. 37.
Remarks. Masure (1991, p. 64) demonstrated that Ato -
podinium has a gonyaulacacean tabulation and that the
archaeopyle of the type is apical, with the formula A1 –4´
,
and with precingular accessory sutures; the operculum is
attached.
Atopodinium cf. haromense Thomas & Cox 1988
(Plate 2, fig. 13)
cf. 1988 Atopodinium haromense Thomas & Cox,
p. 319–321, 323, plate 1, figs 1–6; text-fig. 4.
Age. LO: late Campanian.
Remarks. The Cretaceous forms found in the present
study tend to have slightly more pronounced ornamen-
tation than that of the Jurassic holotype of Atopodinium
haromense (Thomas & Cox 1988, plate 1, figs 1, 2).
Genus Axiodinium Williams, Damassa, Fensome &
Guerstein in Fensome et al. 2009
Type. Williams & Downie 1966b, plate 18, fig. 1, as
Wetzeliella articulata; now Axiodinium prearticulatum.
2009 Axiodinium Williams, Damassa, Fensome &
Guerstein in Fensome et al., p. 16.
Remarks. Both Axiodinium and Apectodinium are wetzeliel-
loidean genera with an equiepeliform archaeopyle (Wil -
liams et al. 2015), but Axiodinium is clearly cavate and
Apectodinium does not have a clear or consistent separation
of the endophragm and periphragm.
Axiodinium augustum (Harland 1979b) Williams
et al. 2015
(Plate 2, figs 14, 15)
1979b Wetzeliella (Apectodinium) augusta Harland,
p. 63, plate 2, figs 13–15.
1981 Apectodinium augustum (Harland) – Lentin &
Williams, p. 14.
2015 Axiodinium augustum (Harland) – Williams et
al., p. 301.
Age. LO: basal Ypresian.
Genus Batiacasphaera Drugg 1970
Type. Drugg 1970, fig. 6A–B, as Batiacasphaera compta.
1970 Batiacasphaera Drugg, p. 813.
Remarks. We adhere to the synopsis presented in Fensome
et al. (2009, p. 17), which specifies that the apical
archaeopyle has an outline lacking or with weakly devel-
oped accessory sutures between precingular plates and a
free operculum.
Batiacasphaera micropapillata Stover 1977
(Plate 2, fig. 16)
1977 Batiacasphaera micropapillata Stover, p. 73,
plate 1, figs 7, 8.
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30
Age. LO (not confirmed in present study) Gelasian on
Scotian Margin (Fensome et al. 2009). Not plotted.
Remarks. This is a species of Batiacasphaera with a gran-
ulate to microreticulate ornamentation.
Genus Batioladinium Brideaux 1975
Type. Alberti 1961, plate 5, fig. 2, as Broomea jaegeri.
1975 Batioladinium Brideaux, p. 124.
1975 Necrobroomea Wiggins, p. 111.
Synopsis. An elongate pareodiniacean cyst, drawn out
into one apical and two generally equal antapical horns.
Acavate. Wall consisting of autophragm. Archaeopyle
apical, with formula A(1–2´), formed from the loss of the
two apical plates.
Remarks. The synopsis is based on the interpretation of
the archaeopyle presented in Wharton (1988, text-fig.
4.13) and Fensome et al. (1993, p. 77, 78). The antapi-
cal horns, though equal on a single specimen, are of
variable length from specimen to specimen.
Batioladinium jaegeri (Alberti 1961) Brideaux
1975
(Plate 2, fig. 17)
1961 Broomea jaergeri Alberti, p. 26, plate 5,
figs 1–7.
1975 Batioladinium jaegeri (Alberti) – Brideaux,
p. 1240.
1975 Necrobroomea jaegeri (Alberti) – Wiggins,
p. 111.
1980 Imbatodinium jaegeri (Alberti) – Dörhöfer &
Davies, p. 37.
1981 Pseudoceratium hansgochtii Lentin &
Williams, p. 236.
Age. LO: Campanian.
Remarks. Most records of this species are from the Early
Cretaceous, but it appears to have an extended range up
to the Campanian in western and northern North
America (e.g. Harker et al. 1990).
Genus Callaiosphaeridium Davey & Williams
1966a
Type. Deflandre & Courteville 1939, plate 4, fig. 1, as
Hystrichosphaeridium asymmetricum.
1966a Callaiosphaeridium Davey & Williams, p. 103.
1967 Hexasphaera Clarke & Verdier, p. 42; name
illegitimate.
Remarks. Callaiosphaeridium is a chorate gonyaulacacean
genus that has large, tubular mesotabular or gonal
processes on the cingulum and may have slender gonal
processes elsewhere. The archaeopyle is epicystal, with
the formula (A1–4´ P1–6´´) and a simple operculum.
Callaiosphaeridium asymmetricum (Deflandre &
Courteville 1939) Davey & Williams 1966a
(Plate 2, fig. 18)
1939 Hystrichosphaeridium asymmetricum Deflandre
& Courteville, p. 100, 101, plate 4, figs 1, 2.
1966a Callaiosphaeridium asymmetricum (Deflandre
& Courteville) – Davey & Williams, p. 104.
1967 Hexasphaera asymmetrica (Deflandre &
Courteville) – Clarke & Verdier, p. 43;
combination illegitimate.
Age. LO: Campanian.
Genus Cannosphaeropsis Wetzel 1933a
Type. Wetzel 1933a, plate 3, fig. 9a, b, as Canno sphae r -
opsis utinensis.
1932 Cannosphaeropsis Wetzel, p. 136; name not
validly published.
1933a Cannosphaeropsis Wetzel, p. 6.
Synopsis. Chorate gonyaulacacean (gonyaulacoidean) cysts
with a subspherical to ellipsoidal central body. Tabulation
on the central body delineated by gonal processes, which
can be restricted to the cingular plates, and sometimes also
intergonal processes, which distally are united by trabecula
that also reflect the tabulation. The gonal processes are
trifurcate distally, the intergonal processes are bifurcate
distally. Archaeopyle precingular, with formula P3´´; oper-
culum free.
Cannosphaeropsis passio de Verteuil & Norris 1996
(Plate 3, fig. 5)
1996 Cannosphaeropsis passio de Verteuil & Norris,
p. 130, 132, 134, 136, plate 7, figs 1–8; plate
8, figs 1–6; plate 17, figs 1, 3–5; text-fig. 33.
Age. LO: Serravallian (based on Piasecki 2003).
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31
Remarks. De Verteuil & Norris (1996) described Canno -
sphaeropsis passio as having cingular processes only, even
though the trabecula delineate the complete tabulation.
Genus Cerebrocysta Bujak in Bujak et al. 1980
Type. Bujak et al. 1980, plate 13, figs 4–5, as Cere bro -
cysta bartonensis.
1980 Cerebrocysta Bujak in Bujak et al., p. 42.
Remarks. The synopsis for Cerebrocysta provided by Fen -
some et al. (2009, p. 18) covers all the salient details of the
morphology of this genus. These authors also noted that
it is difficult to separate Cerebrocysta from Pyxidinopsis.
The only distinctions are in the possible variability in the
number of precingular plates lost in archaeopyle forma-
tion in Cerebrocysta and perhaps different wall structures.
Regardless, we follow Fensome et al. (2009) in assigning
Cenozoic taxa with the general morphology of Cere -
brocysta and Pyxidinopsis to Cerebrocysta.
Cerebrocysta bartonensis Bujak in Bujak et al. 1980
(Plate 2, figs 19, 20)
1980 Cerebrocysta bartonensis Bujak in Bujak et al.,
p. 42, plate 13, figs 4–7.
Age. LO: Bartonian.
Cerebrocysta magna Bujak 1994
(Plate 3, fig. 6)
1994 Cerebrocysta magna Bujak, p. 121, plate 2,
figs 10, 11.
Age. LO: Lutetian.
Genus Cerodinium Vozzhennikova 1963
Type. Vozzhennikova 1963, text-fig. 9, as Cerodinium
sibiricum.
1963 Cerodinium Vozzhennikova, p. 181.
1963 Ceratiopsis Vozzhennikova, p. 181; name ille-
gitimate.
Remarks. Fensome et al. (2009, p. 18) noted that Cero -
dinium is characterised by having an isodeltaform to
isothetaform 2a plate, a free perioperculum and a
symmetrical antapex. The endoarchaeopyle may involve
all three anterior intercalary plates. At present it does not
seem practicable to separate into distinct species forms
that lose the 2a plate only from those that lose the 1a, 2a
and 3a plates individually from the endocyst. However,
such a separation seems to have some stratigraphic value
as forms occurring in younger rocks all seem to lose the
2a plate only; earlier forms may lose all three anterior
intercalaries.
Cerodinium diebelii (Alberti 1959) Lentin &
Williams 1987
(Plate 3, figs 1–4)
1959 Deflandrea diebelii Alberti, p. 99, 100, plate 9,
figs 18–21.
1967 Ceratiopsis diebelii (Alberti) – Vozzhennikova,
p. 159; combination illegitimate.
1987 Cerodinium diebelii (Alberti) – Lentin &
Williams, p. 114.
Age. LO: latest Danian.
Cerodinium glabrum (Gocht 1969) Fensome et al.
2009
(Plate 3, figs 7, 8)
1969 Deflandrea speciosa forma glabra Gocht, p. 10,
text-fig. 3.
1973 Deflandrea speciosa subsp. glabra (Gocht) –
Lentin & Williams, p. 45.
1977a Ceratiopsis speciosa subsp. glabra (Gocht) –
Lentin & Williams, p. 21 (combination ille-
gitimate).
1987 Cerodinium speciosum subsp. glabrum (Gocht)
– Lentin & Williams, p. 115.
2009 Cerodinium glabrum (Gocht) – Fensome et al,
p. 19.
Age. LO: latest Thanetian (Late Paleocene).
Cerodinium kangiliense Nøhr-Hansen &
Heilmann-Clausen 2001
(Plate 3, figs 9, 10)
2001 Nøhr-Hansen & Heilmann-Clausen, p. 158,
160, 162–164, fig. 4, nos 1–9, fig. 5, nos 1–9.
Age. LO: Selandian.
Cerodinium speciosum (Alberti 1959) Lentin &
Williams 1987
(Plate 3, fig. 13)
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32
1959 Deflandrea speciosa Alberti, p. 97, plate 9,
figs 12, 13.
1977a Ceratiopsis speciosa (Alberti) – Lentin &
Williams, p. 21; combination illegitimate.
1987 Cerodinium speciosum (Alberti) – Lentin
&Williams, p. 115.
Age. LO: latest Thanetian.
Cerodinium striatum (Drugg 1967) Lentin &
Williams 1987
(Plate 3, figs 11, 12)
1967 Deflandrea striata Drugg, p. 18, plate 2,
figs 13, 14.
1977a Ceratiopsis striata (Drugg) – Lentin &
Williams, p. 21; combination illegitimate.
1987 Cerodinium striatum (Drugg) – Lentin &
Williams, p. 115.
Age. LO Thanetian. Not plotted.
Remarks. Cerodinium striatum has a distinct develop-
ment of more or less parallel folds. The folds should not
be confused with the plications of Cerodinium diebelii
(which is distinctly elongate) or the linearly aligned
granules or denticles in Cerodinium speciosum.
Genus Charlesdowniea Lentin & Vozzhennikova
1989 emend. Williams et al. 2015
Type. Williams & Downie 1966b, plate 18, fig. 8, text-
fig. 47, as Wetzeliella coleothrypta, and Bujak et al. 1980,
plate 12, figs 7, 8, as Kisselevia coleothrypta.
1989 Charlesdowniea Lentin & Vozzhennikova, p.
225, 227.
Remarks. The emended diagnosis of Williams et al.
(2015) is followed; this emphasises the archaeopyle type
and the nature of the pericystal ornamentation. The
archaeopyle of Charlesdowniea is equiepeliform and the
processes are intratabular and sometimes also penitabu-
lar. Distally, the processes on individual plates are unit-
ed by membranes.
Charlesdowniea coleothrypta (Williams & Downie
1966b) Lentin & Vozzhennikova 1989
(Plate 4, fig. 9)
1966b Wetzeliella coleothrypta Williams & Downie,
p. 185, 186, plate 18, figs 8, 9; text-fig. 47.
1976 Kisselevia coleothrypta (Williams & Downie) –
Lentin & Williams, p. 136.
1989 Charlesdowniea coleothrypta (Williams &
Downie) – Lentin & Vozzhennikova, p. 225.
Age. LO: Ypresian. Not plotted.
Remarks. In the Labrador Margin samples, Charles dow -
n iea coleothrypta has its LO just below that of Scale no -
dinium scalenum, but is not common.
Genus Chatangiella Vozzhennikova 1967 emend.
nov.
Type. Vozzhennikova 1967, plate 56, fig. 1; plate 57,
fig. 1, as Chatangiella niiga.
1967 Chatangiella Vozzhennikova, p. 128, 129.
1967 Australiella Vozzhennikova, p. 129, 130.
1967 Cooksoniella Vozzhennikova, p. 183, 184.
Emended description. Peridiniacean (deflandreoid) cysts
that are proximate, dorsoventrally compressed and peri-
dinioid in outline, with epicystal ‘shoulders’ and asym-
metrical antapex, the left side being larger. Cir cumcavate
to, generally, bicavate. Surface generally atabulate to
weakly paratabulate, but cingulum clearly reflected by
ridges, commonly serrated, reflecting the positions of
pre- and postcingular plates. Periarchaeopyle intercalary,
with formula I2a, operculum free or attached; plate 2a is
lati- to iso-omegaform hexa; endoarchaeopyle with for -
mula I2a or I1a +2a +3a.
Remarks. Fensome et al. (2009, p. 19) provided a synop-
sis for Chatangiella that encompasses all its salient morpho-
logic features. As part of their synopsis, these authors
stated: “Periarchaeopyle intercalary, with formula I2a,
operculum free or attached; plate 2a is isodelta form to
(typically) iso-omegaform hexa; endoarchaeopyle with
formula I2a or I1a + 2a + 3a.” We disagree with the synopsis
of Fensome et al. (2009) in two important respects.
Firstly, only taxa with omegaform 2a plates should be
included in Chatangiella; and, secondly, taxa with 2a
plates that are lati-omegaform must be included since the
type of Chatangiella (Vozzhennikova 1967, plate 56, fig.
1; plate 57, fig. 1) has such an archaeopyle. Thus it is
proposed here that a characteristic feature of Chatangiella
be the possession of a lati- or iso-omegaform hexa 2a
plate.
In their emendation of Chatangiella, Lentin & Wil -
liams (1976, p. 51, 52) emphasised the omegaform shape
of the pericystal 2a plate and its common attachment
Bulletin36.qxp_Bulletin 36 19/12/16 13.39 Side 32
33
posteriorly. Unfortunately, the critical shape and rela-
tionship of this plate have been overlooked in many
subsequent papers. To correct this concern, the omega -
form shape of the operculum is re-emphasised in the
emended description above. Although epicystal shoul-
ders, a distinctive characteristic of this genus, appear to
almost always be related to the presence of an omegaform
periarchaeopyle, which feature came first is open to
debate.
As noted under Alterbidinium, the morphological
differences used to separate the deflandreoid genera are
in large part unsatisfactory. This criticism applies to
Chatangiella: to bring stability to the genus, we propose
that any peridiniacean forms with a partite cingulum
that do not have an omegaform periarchaeopyle should
be re-assigned.
Chatangiella decorosa (McIntyre 1975) Lentin &
Williams 1976
1975 Deflandrea decorosa McIntyre, p. 63, 64, plate
2, figs 1–4.
1976 Chatangiella decorosa (McIntyre) – Lentin &
Williams, p. 54.
Age. LO: early Campanian.
Remarks. McIntyre (1975, p. 64) noted that Chatangiella
decorosa differs from Chatangiella ditissima in being
larger. The size of Chatangiella ditissima overlaps that of
Chatangiella tripartita. Chatangiella decorosa also has
more pustules than Chatangiella ditissima.
Chatangiella madura Lentin & Williams 1976
(Plate 3, figs 14, 15)
1970b Deflandrea manumii Cookson & Eisenack,
p. 141, 142, plate 11, figs 10, 11.
1976 Chatangiella manumii (Cookson & Eisenack)
– Lentin & Williams, p. 54; name illegitimate.
1976 Chatangiella madura (Cookson & Eisenack) –
Lentin & Williams, p. 54.
Age. LO: Campanian.
Chatangiella tripartita (Cookson & Eisenack
1960a) Lentin & Williams 1976
(Plate 3, figs 16, 20)
1960a Deflandrea tripartita Cookson & Eisenack,
p. 2, 3, plate 1, fig. 10.
1967 Australiella tripartita (Cookson & Eisenack) –
Vozzhennikova, p. 134, 135.
1976 Chatangiella tripartita (Cookson & Eisenack)
– Lentin & Williams, p. 55.
Age. LO: Campanian (Late Cretaceous). Not plotted.
Remarks. There is considerable confusion concerning
the morphology of Chatangiella tripartita and the rela-
ted form Chatangiella victoriensis. Both are surprisingly
large. According to Cookson & Eisenack (1960a, p. 3),
the pericyst of Chatangiella tripartita ranges from 100
to 120 μm in length and 59 to 71 μm in width.
Cookson & Manum (1964) stated that in Chatangiella
victoriensis, the pericyst was 76 to 116 μm in length and
49–73 μm in width. Comparison of the sizes of the
above two species with one of the higher latitude
(Arctic) species of Chatangiella, Chatangiella ditissima,
shows that there is some overlap, especially in width.
McIntyre (1975, p. 63) noted that the pericyst in
Chatangiella ditissima could be 115–150 μm long and
60–90 μm wide. The similar but even larger, higher
latitude species Chatangiella decorosa has a pericyst that
can be 130–175 μm long and 80–110 μm wide.
Our conclusion from comparison of several Chatan -
giella species is that size is not a reliable characteristic
when separating species. But size has some significance:
for example specimens of Chatangiella found on the
Grand Banks of Newfoundland are generally much
smaller than those from the high Arctic. Gigantism is a
common feature of life in higher latitudes, so it is no
surprise to find it among dinoflagellates.
Differences in the expression of the cingulum and the
surface ornamentation of the pericyst are perhaps more
diagnostic. Cookson & Manum (1964, p. 521, 522)
noted that the cingulum of the holotype of Chatangiella
tripartita is denoted by two pairs of ill-defined short,
offset, low parallel ridges on the ventral surface and a
fold-like line on the dorsal surface. The ridges are not
serrate or denticulate, a feature of many specimens placed
in Chatangiella tripartita. Regarding the cingulum of
Chatangiella victoriensis, Cookson & Manum (1964)
stated that it “is bordered by conspicuous ridges or by
linearly arranged wart-like thickenings of varying size
and shape”. McIntyre (1975, p. 62, 63) described
Chatangiella ditissima as having a cingulum delineated by
“discontinuous ridges consisting of rows of pustules that
may join to form narrow grooves”. That author recog-
nised seven and five ridges on the anterior and posterior
margins respectively of the cingulum, which represent
the difference in number of plates between the pre- and
Bulletin36.qxp_Bulletin 36 19/12/16 13.39 Side 33
34
postcingular series. These two series of plates can also
have penitabular pustules mirroring the plate outlines.
Cookson & Manum (1964, p. 522) gave a thickness
of 1.0–1.7 μm for the periphragm (as the thecal wall) of
Chatangiella victoriensis and noted that it is “ornamen-
ted with fairly evenly scattered rod-like projections c.
0.5–1.5 μm long; in surface view the rods appear as dots
usually between 0.5 and 1.0 μm in diameter but a few
smaller and larger ones are usually present”. In Cha -
tangiella tripartita, the periphragm is finely granular.
McIntyre (1975) described the periphragm of Cha -
tangiella ditissima as smooth, except for the pustules
delineating the tabulation and occasional pits.
What is the conclusion to be drawn from the above
and from our observations on the specimens from the
offshore wells? The most compelling is the extreme
variability in the size of the taxa and the degree of
expressions of tabulation. In this paper, all the ‘smaller’
forms are assigned to Chatangiella tripartita if they lack
pustules other than on the cingulum. We would like to
include those smaller forms with pustules defining the
pre- and postcingular plates in a new species but do not
have enough specimens to do that here.
Genus Chiropteridium Gocht 1960 emend. nov.
Type. Gocht 1960, plate 17, fig. 1, as Chiropteridium
lobospinosum.
1959 Galea Maier, p. 305; name illegitimate.
1960 Chiropteridium Gocht, p. 221.
Emended diagnosis. Areoligeracean cysts that are proxi-
mochorate to chorate, with subrounded to lenticular
central body that may be asymmetrical antapically.
Cavate or acavate; if cavate, with cavation mainly
restricted to scalloped marginal wings; when acavate
there may be marginal wings and/or processes. Mid-
ventral and mid-dorsal areas devoid of processes or
having processes of a reduced size. Surface atabulate or
with intratabular processes. Archaeopyle apical, with
formula A(1–4´) and free operculum; sulcal notch offset
to the left.
Remarks. Chiropteridium has not been emended previ-
ously. However, Stover & Evitt (1978) provided a
synopsis and modified description and Fensome et al.
(2009, p. 21) gave a synopsis. In their synopsis of
Chiropteridium, Stover & Evitt (1978, p. 27) stated:
“Cysts skolochorate, body lenticular; processes isolated
or connected proximally and absent or greatly reduced
in size and number on ventral surface; archaeopyle
apical, Type tA; parasulcal notch offset.” In their modi-
fied description, the same authors noted that processes
could be isolated or partly connected in longitudinal
rows, with the connections being proximal. Also they
stated that processes are absent or reduced in size and
numbers ventrally. Fensome et al. (2009, p. 21) were
more specific in providing the following synopsis for
Chiropteridium: “Areoligeracean cysts that are proximo-
chorate to chorate, with lenticular central body that
may be asymmetrical antapically. Cavate or acavate; if
cavate, with cavation mainly restricted marginally to
‘wings’ and scalloped; if acavate, marginate wings
formed from longitudinal crests or from longitudinally
taeniate processes. Surface atabulate. Archaeopyle
apical, with formula A(1–4´), operculum free; sulcal
notch offset to the left.”
Fensome et al. (2009, p. 21) stated that “Chirop -
teridium is characterised by lateral extensions or ‘wings’
that may be cavate; these wings are typically scalloped to
varying degrees or formed by taeniate processes. In
Membranophoridium, processes are absent and the ‘wings’
are continuous pericoelar sacs.” Although correct for
species included to date in Chiropteridium, forms are
recorded here that possess only processes that are not
connected proximally or along their length and which
denote the tabulation. These are accommodated in the
above emended diagnosis for Chiropteridium.
Chiropteridium galea (Maier 1959) Sarjeant 1983
(Plate 3, figs 17–19)
1959 Galea galea Maier, p. 306, plate 29, fig. 4;
text-fig. 2.
1959 Galea levis Maier, p. 308, plate 30, figs 1, 2.
1959 Galea mespilana Maier, p. 306, 307, plate 29,
figs 5, 6.
1960 Chiropteridium dispersum Gocht, p. 227, plate
18, figs 1–16; text-figs 16–27.
1961 Membranophoridium multispinatum Gerlach,
p. 203, 204, plate 29, fig. 5.
1961 Membranophoridium partispinatum Gerlach,
p. 201, plate 29, fig. 6.
1963 Chiropteridium partispinatum (Gerlach) –
Brosius, p. 48.
1964 Baltisphaeridium leve (Maier) – Sarjeant, p. 176.
1964 Baltisphaeridium mespilanum (Maier) –
Sarjeant, p. 176.
1969 Cleistosphaeridium leve (Maier) – Davey et al.,
p. 16.
1973 Chiropteridium mespilanum (Maier) – Lentin
& Williams, p. 26.
Bulletin36.qxp_Bulletin 36 19/12/16 13.39 Side 34
35
1975 Hystrichosphaeridium mespilanum (Maier) –
Eisenack & Kjellström, p. 233, 234.
1983 Chiropteridium galea (Maier) – Sarjeant,
p. 108.
Age. LO: Chattian.
Chiropteridium gilbertii sp. nov.
(Plate 4, figs 1–4)
Holotype. Plate 4, fig. 4, from a cuttings sample at
1600–1610 m in Gilbert F-53, GSC type collection
no.137976, sample P39466, slide 01, co-ordinates 17.4
× 106.0, England Finder R36/3. Overall length 83 μm,
width 87 μm; central body, length 57 μm, width 52
μm; maximum length of processes 20 μm, width varies
from less than 1 to 10 μm. The age determined for the
sample from which the holotype was recovered is basal
Bartonian.
Etymology. The epithet is derived from the name of the
Gilbert F-53 offshore exploration well, in which the
species is abundant.
Diagnosis. A species of Chiropteridium in which the
processes or process complexes delineate the tabulation
and there are no interconnections between processes
representing adjacent plates.
Description. The processes delineate the tabulation,
though none are interconnected. Processes on the mid-
ventral and mid-dorsal surfaces are reduced in width or
absent: this is especially true of the 3´´ process that, if
present, is very slender. Commonly, processes are basal-
ly trumpet-shaped, then tubular or parallel-sided. Pro -
cesses are also of variable size depending on which plate
series they are reflecting, with cingular processes being
invariably slender, precingular processes broader, and
postcingular processes being broader still, and the
antapical process being broadest of all. Many of the
processes are perforate.
Size. Overall maximum diameter 87 μm; maximum
diameter of central body 57 μm; maximum length of
processes 25 μm, maximum width 17 μm; seven speci-
mens measured.
Age. LO: Bartonian.
Remarks. In Chiropteridium gilbertii, the central body,
which is granulate, commonly has a prominent antapi-
cal protuberance. All of the processes, which are in -
tratabular, have a circular cross-section proximally and
distally are closed and commonly branch into short
bifurcations or aculeae. Proximally, the processes tend
to be conical then become tubular about halfway along
their length before flaring distally. The processes are
often fibrous and/or perforate, especially the antapical
process, which sometimes subdivides to form a process
complex. Recognition of the antapical process, if the
archaeopyle is not obvious, is facilitated by the extreme
width and the perforations, which may form arches in
the process wall. Cingular processes are invariably slen-
der, some less than 1 μm wide, rarely exceed four in
number, and are restricted to the ambital region of the
central body. Apical processes show variation in width
on different specimens. Precingular processes are always
narrower than the postcingular processes. Although the
sulcal notch is offset, this is not always obvious.
Chiropteridium gilbertii differs from Chirop te ri dium
galea in having processes or process complexes that are
restricted to individual plates. Licracysta semicirculata
has processes that are restricted to the ambitus and tend
to form arcuate complexes. The processes of Chiro p -
teridium gilbertii vary in size according to the plate they
represent, and are often fibrous and/or perforate.
Chirop teridium conispinum also has membranous pro -
cesses, but these are restricted to the dorsal surface, with
two linear membranes running apically–antapically on
the ventral surface.
Genus Chlamydophorella Cookson & Eisenack
1958
Type. Cookson & Eisenack 1958, plate 11, fig. 1, as
Chlamydophorella nyei.
1958 Chlamydophorella Cookson & Eisenack, p. 56.
Remarks. As Fensome et al. (2009, p. 21) noted in their
synopsis for Chlamydophorella, the genus is characterised
by being proximate, holocavate, with short, solid
processes forming buttresses between the autophragm
and ectophragm, an apical horn formed from the
ectophragm, and an apical archaeopyle. The tabulation
is gonyaulacacean, but whether it shows neutral or
dextral torsion is not known. Sepispinula differs from
Chlamydophorella in lacking an ectophragm.
Chlamydophorella nyei Cookson & Eisenack 1958
(Plate 4, fig. 5)
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36
1958 Chlamydophorella nyei Cookson & Eisenack,
p. 56, plate 11, figs 1–3.
1970b Chlamydophorella apiculata Cookson &
Eisenack, p. 150, 151, plate 13, fig. 3.
1970b Chlamydophorella lagena Cookson &
Eisenack, p. 151, plate 13, fig. 4.
Age. LO: Coniacian (Late Cretaceous).
Chlamydophorella cf. nyei Cookson & Eisenack
1958
(Plate 4, figs 6–8)
Age. LO : Campanian. Not plotted.
Remarks. This form includes specimens similar to Chla -
mydophorella nyei but which lack an evident apical
horn.
Genus Chytroeisphaeridia (Sarjeant 1962) Downie
& Sarjeant 1965
Type. Sarjeant 1962, plate 70, fig. 13, as Leiosphaeridia
subgenus Chytroeisphaeridia chytroeides.
1962 Leiosphaeridia subgenus Chytroeisphaeridia
Sarjeant, p. 492.
1965 Chytroeisphaeridia Downie & Sarjeant, p. 102.
Chytroeisphaeridia hadra sp. nov.
(Plate 4, figs 17, 18)
Holotype. Plate 4, fig. 18 from a cuttings sample at
3120–3140 m in Roberval K-92, GSC type collection
no. 137902, sample P17728, slide 01, co-ordinates
19.5 × 109.8, England Finder U40/1. Overall length
79 μm, width 81 μm; wall thickness 5 μm, archaeopyle
length 37 μm, width 33 μm. The age determined for
the sample from which the holotype was recovered is
Late Cretaceous (the specimen is caved).
Etymology. The epithet is from the Greek hadros mean-
ing well-developed, bulky, stout, strong, great, in refer-
ence to the thick, sturdy wall.
Diagnosis. A species of Chytroeisphaeridia with a thick
wall that has a subdued rugulate, sometimes lightly stri-
ated surface. The archaeopyle is large, apparently result-
ing from the loss of the 3´´ plate only.
Size. Overall length 69–82 μm; width 64–81 μm. Six
specimens measured.
Age. LO: Bartonian.
Remarks. Some specimens of Chytroeisphaeridia hadra
have parallel striations on the wall but these are irregular
and discontinuous. The archaeopyle is large, suggesting
that more than a single plate may be involved.
Genus Cleistosphaeridium Davey et al. 1966
Type. Davey et al. 1966, plate 10, fig. 7, as Cleisto -
sphaeridium diversispinosum.
1966 Cleistosphaeridium Davey et al., p. 166.
Remarks. We concur with the retention and emenda-
tion of Cleistosphaeridium as proposed by Eaton et al.
(2001, p. 176, 177) and the synopsis provided by
Fensome et al. (2009, p. 22). Following Fensome et al.
(2007, p. 408), Cleistosphaeridium is considered to be
areoligeracean because of the asymmetry in some spe -
cimens of the antapex and sulcal notch, and also
because it forms a morphological plexus with other
areoligeracean genera, including Glaphyrocysta, Enne -
adocysta, Licracysta and Cooksonidium. We include in
the genus species in which adjacent processes are medi-
ally and distally interconnected.
Cleistosphaeridium diversispinosum Davey et al.
1966
(Plate 4, figs 19, 20)
1966 Cleistosphaeridium diversispinosum Davey et
al., p. 167, plate 10, fig. 7.
1993 Systematophora diversispinosa (Davey et al.) –
Islam, p. 88.
Age. LO: Serravallian.
Remarks. Following Fensome et al. (2009, p. 22), Cleisto -
sphaeridium diversispinosum and Cleistosphaeridium ancyr -
eum are not considered synonyms.
Cleistosphaeridium elegantulum sp. nov.
(Plate 4, figs 10–12)
Holotype. Plate 4, fig. 12, from a cuttings sample at
2286.03 to 2295.17 m in Karlsefni A-13, GSC type
collection no. 138033, sample P39600, slide 01, co-ordi-
nates 17.9 × 101.7, England Finder S44.2. Central body
maximum diameter 50 μm, maximum length of process-
es about 25 μm. The age determined for the sample from
which the holotype was recovered is Lutetian–Bartonian.
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37
Etymology. From the Latin elegantulus (very fine), in
reference to the narrow, long processes.
Diagnosis. A species of Cleistosphaeridium with nume -
rous long, fine, flexible, unconnected processes, some
to many of which are dolabrate distally. The length of
many of the processes equals about one half of the
central body diameter. Proximal ridges are absent to
weakly developed.
Size. Central body diameter 30‒57 μm; maximum
length of processes up to half maximum diameter of
central body; three specimens measured.
Age. LO: Lutetian‒Bartonian. Not plotted.
Remarks. This species differs from other species of
Cleistosphaeridium in its long, fine, flexible processes.
Cleistosphaeridium palmatum sp. nov.
(Plate 4, figs 13–16)
Holotype. Plate 4, figs 13, 14 from a cuttings sample at
2450–2460 m in Roberval K-92: GSC type collection
no. 137897, sample P17706, slide 01, co-ordinates 6.9
× 100.6, England Finder G30/0‒2. Maximum overall
diameter 87 μm, maximum diameter of central body 50
μm, maximum length of processes 25 μm. The age
determined for the sample from which the holotype
was recovered is early Ypresian.
Etymology. The epithet is from the Latin palmatus,
meaning marked or shaped like the palm of the hand,
in reference to the shape of the distal terminations of
the processes.
Diagnosis. A species of Cleistosphaeridium in which the
solid processes are of irregular width, varying from 1–7
μm and distally slender to splayed, with some adjacent
processes interconnected distally, commonly so that
they form arches.
Description. The processes are of irregular width, distal-
ly slender to splayed, with some adjacent processes
interconnected distally. They are predominantly dolo-
brate, but some are bifid. The wall of the central body
is generally less than 1 μm thick. Archaeopyle apical,
with formula A(1–4´), operculum attached or free. Other
aspects of the tabulation cannot be determined, as spec-
imens possess more than one process per plate, some-
times in complexes. Where present, the complexes appear
to occur on the precingular, postcingular and antapical
plates.
Size. Maximum diameter of central body 48 μm, length
of processes 10–25 μm, four specimens measured.
Age. LO: late Ypresian.
Remarks. The processes of specimens belonging to this
species show similarities to those found in Adnato -
sphaeridium, Enneadocysta, Licracysta and other species of
Cleistosphaeridium. Although the distal branches of pro -
cesses can sometimes be interconnected, the general
absence of trabecula uniting processes precludes assign-
ment of the species to Adnatosphaeridium. Cleistospha er -
idium palmatum differs from species of Enneadocysta
because the number of processes per plate (where dis -
cernible as being related to tabulation) invariably exceeds
one, and distally processes are dolobrate rather than
licrate or clypeate. Moreover, the antapex of Cleis to s phae r -
idium palmatum is not characterised by two pro cesses
(see Fensome et al. 2007, p. 394). Cleistospha eridium
diversispinosum differs in not having branched processes
in which the branches often meet along their length or
distally and in not having spatulate endings. Specimens
of the species Cleistosphaeridium polypetellum are slender
and are not spatulate or branched distally; species of
Licracysta have proximal and ventral surfaces on which
processes are absent or reduced.
Cleistosphaeridium polypetellum (Islam 1983b)
Stover & Williams 1995
(Plate 5, figs 1, 2)
1983b Areosphaeridium polypetellum Islam, p. 82, 84,
plate 2, figs 1–6.
1995 Cleistosphaeridium polypetellum (Islam) –
Stover & Williams, p. 102.
Age. LO: Ypresian.
Remarks. Some connections occur between adjacent
processes, differentiating this species from Cleistos phae r -
idium diversispinosum and Cleistosphaeridium an cyreum.
Processes in Cleistosphaeridium polypetellum are less
variable and the species lacks the thick processes of Clei -
sto s phaeridium palmatum. Distally, most processes of
Cleisto sphaer idium polypetellum are strongly dola brate
and sometimes licrate. However, a few simple bifid
processes occur.
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38
Genus Cordosphaeridium Eisenack 1963a
Type. Klumpp 1953, plate 18, figs 1, 2, as Hystricho -
sphaeridium inodes.
1963a Cordosphaeridium Eisenack, p. 261.
1981 Tityrosphaeridium Sarjeant, p. 120.
Remarks. We concur with the synopsis for Cordos phae r -
idium provided by Fensome et al. (2009, p. 22). The
characteristic features of the genus are the spheroidal
central body, the precingular (P3´´) archaeopyle and the
fibrous wall and processes. The processes, which are
commonly cylindrical and restricted to one per plate,
are generally of approximately equal length but can
show considerable variation in width.
Cordosphaeridium cantharellus (Brosius 1963)
Gocht 1969
(Plate 5, fig. 7)
1963 Hystrichosphaeridium cantharellus Brosius, p.
40, 41, plate 6, fig. 1; text-fig. 2, nos 11a–c.
1969 Cordosphaeridium cantharellus (Brosius) –
Gocht, p. 45.
1981 Tityrosphaeridium cantharellus (Brosius) –
Sarjeant, p. 120.
Age. LO: Burdigalian.
Remarks. As in the holotype, the process width in the
Labrador Margin specimens of Cordosphaeridium can -
tharellus shows considerable variation. The species was
retained in Cordosphaeridium by Edwards (2001, p. G19).
Cordosphaeridium delimurum Fensome et al. 2009
(Plate 5, figs 3, 4)
2009 Cordosphaeridium delimurum Fensome et al.,
p. 23, plate 2, figs l–q.
Age. LO: early Lutetian. Not plotted.
Remarks. As its name implies, Cordosphaeridium deli-
murum differs from Cordosphaeridium inodes and Cor -
dosphaeridium gracile in having a much thinner central
body wall. Unlike Cordosphaeridium fibrospinosum, Cor -
do sphaeridium delimurum has solid rather than perfo -
rate process walls.
Cordosphaeridium fibrospinosum Davey &
Williams 1966a
(Plate 5, figs 5, 6)
1966a Cordosphaeridium fibrospinosum Davey &
Williams, p. 86, plate 5, fig. 5.
1966a Cordosphaeridium exilimurum Davey &
Williams, p. 87, 88, plate 11, fig. 2.
1970 Achomosphaera valianta Sah et al., p. 145,
plate 1, figs 8, 9.
1978 Cordosphaeridium valiantum (Sah et al.) –
Stover & Evitt, p. 147.
1981 Hystrichosphaerina? exilimura (Davey
&Williams) – Sarjeant, p. 122.
1981 Emmetrocysta? fibrospinosa (Davey &
Williams) – Sarjeant, p. 123.
1986 Tityrosphaeridium? exilimurum (Davey &
Williams) – Jain & Garg, p. 120.
1986 Tityrosphaeridium? fibrospinosum (Davey &
Williams) – Jain & Garg, p. 121.
Age. LO: late Rupelian. Not plotted.
Remarks. As noted above, Cordosphaeridium fibrospino -
sum has processes with fibrous, perforate walls. We agree
with Fensome et al. (2009) that Cordosphaeridium exil-
imurum and its long-recognised synonym Achomo -
sphaera valianta cannot be meaningfully differentiated
from Cordosphaeridium fibrospinosum. Although this spe -
cies has an Oligocene LO, at least in offshore eastern
Canada, it tends to be most common in the Paleocene.
Cordosphaeridium funiculatum Morgenroth 1966a
(Plate 5, fig. 8)
1966a Cordosphaeridium funiculatum Morgenroth,
p. 22, 23, plate 6, figs 2–3.
1981 Tityrosphaeridium funiculatum (Morgenroth)
– Sarjeant, p. 121.
Age. LO: Priabonian.
Cordosphaeridium gracile (Eisenack 1954) Davey
& Williams 1966a
(Plate 5, fig. 9)
1954 Hystrichosphaeridium inodes subsp. gracile
Eisenack, p. 66, plate 8, fig. 17; plate 10,
figs 3–8; plate 12, figs 7, 21.
1963a Cordosphaeridium inodes subsp. gracile
(Eisenack) – Eisenack, p. 261.
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39
1966a Cordosphaeridium gracile (Eisenack) – Davey
& Williams, p. 84.
1981 Tityrosphaeridium gracile (Eisenack) –
Sarjeant, p. 121.
Age. LO: early Lutetian.
Cordosphaeridium inodes (Klumpp 1953) Eisenack
1963a
(Plate 5, fig. 10)
1953 Hystrichosphaeridium inodes Klumpp, p. 391,
plate 18, figs 1, 2.
1963a Cordosphaeridium inodes (Klumpp) –
Eisenack, p. 261.
Age. LO: consistent occurrence early Lutetian. Not
plotted.
Remarks. The holotype of Cordosphaeridium inodes
(Klumpp 1953, plate 18, figs 1, 2) shows considerable
variation in the width of the processes, some being
remarkably similar to those of Cordosphaeridium gracile.
It is this variability in width, however, that is the distin-
guishing characteristic of Cordosphaeridium inodes.
Genus Cribroperidinium Neale & Sarjeant 1962
Type. Neale & Sarjeant 1962, plate 19, fig. 4, text-fig.
3a, b, as Cribroperidinium sepimentum.
1962 Cribroperidinium Neale & Sarjeant, p. 443.
1968 Acanthaulax Sarjeant, p. 227.
1978 Millioudodinium Stover & Evitt, p. 173.
1984b Meristaulax Sarjeant, p. 160.
1988 Meristaulax Brenner, p. 65.
Synopsis. Gonyaulacacean (cribroperidinioid) cysts that
are proximate, with spheroidal to more commonly
ovoidal central body, usually surmounted by an apical
horn. Acavate or cornucavate. Tabulation strongly delin-
eated by sutural and penitabular ornamentation. Archae -
o pyle pre cingular, with formula P3´´, operculum free.
Cribroperidinium giuseppei (Morgenroth 1966a)
Helenes 1984
1966a Gonyaulax giuseppei Morgenroth: 5, plate 2,
figs 3–6.
1969 Gonyaulacysta giuseppei (Morgenroth) –
Sarjeant, p. 9.
1978 Millioudodinium? giuseppei (Morgenroth) –
Stover & Evitt, p. 174.
1982 Rhynchodiniopsis? giuseppei (Morgenroth) –
Sarjeant, p. 36.
1984 Cribroperidinium giuseppei (Morgenroth) –
Helenes, p. 121.
Age. Local acme: Priabonian.
Remarks. Cribroperidinium giuseppei is a species of Cribro -
peridinium with a small, button-like apical horn, a spon -
gy wall, distinctly reflected paratabulation and a low,
coarse reticulum superimposed on the paraplates. This
includes many forms assigned in the literature to Cri -
broperidinium tenuitabulatum, but the latter species is
now considered to be a junior synonym of Apteodinium
australiense. Helenes (1984, p. 122) noted that the sutur-
al ridges in Cribroperidinium giuseppei are smooth and
that the penitabular ridges are more commonly found on
the hypocyst.
Genus Cyclonephelium Deflandre & Cookson 1955
Type. Deflandre & Cookson 1955, plate 2, fig. 12, as
Cyclonephelium compactum.
1955 Cyclonephelium Deflandre & Cookson,
p. 285.
1961 Circulodinium Alberti, p. 28.
Remarks. We follow the synopsis for this genus pro -
posed by Fensome et al. (2009, p. 24), and agree with
those authors in considering Circulodinium to be a
junior synonym of Cyclonephelium.
Cyclonephelium distinctum (Deflandre & Cookson
1955) Jansonius 1986
(Plate 6, fig. 10)
1955 Cyclonephelium distinctum Deflandre &
Cookson, p. 285, 286, plate 2, fig. 14; text-
figs 47, 48.
1961 Circulodinium hirtellum Alberti, p. 28, 29,
plate 4, fig. 20.
1969 Canningia hirtella (Alberti) – Millioud,
p. 425.
1978 Cyclonephelium hirtellum (Alberti) – Davey,
p. 894.
1986 Circulodinium distinctum (Deflandre &
Cookson) – Jansonius, p. 204.
Age. LO: latest Maastrichtian.
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40
Genus Dapsilidinium Bujak et al. 1980
Type. Davey & Williams 1966a, plate 4, fig. 10, as Poly -
sphaeridium pastielsii.
1980 Dapsilidinium Bujak et al., p. 27, 28.
Remarks. For this genus, the concept of Fensome et al.
(2009, p. 26) is followed.
Dapsilidinium pseudocolligerum (Stover 1977)
Bujak et al. 1980
(Plate 5, fig. 13)
1977 Polysphaeridium pseudocolligerum Stover, p.
74, 75, plate 1, figs 14–19.
1980 Dapsilidinium pseudocolligerum (Stover) –
Bujak et al., p. 28.
Age. LO: Tortonian.
Remarks. As Fensome et al. (2009) observed, Dapsili -
dinium pastielsii and Dapsilidinium pseudocolligerum differ
primarily in the proximal morphology of the processes. In
Dapsilidinium pastielsii, the processes are initially broad
before tapering gradually, whereas in Dapsilidinium pseu -
docolligerum they are more or less cylindrical proximally,
tapering gradually to the distal opening.
Dapsilidinium pseudoinsertum sp. nov.
(Plate 5, figs 11, 12)
Holotype. Plate 5, fig. 12 from a cuttings sample at
2825‒2835 m in Rut H-11, GSC type collection no.
137957, sample P39388, slide 01, co-ordinates 19.6 ×
105.1, England Finder U48/1‒2. Central body maxi-
mum diameter 32 μm; length of processes up to about 15
μm. The age determined for the sample from which the
holotype was recovered is Ypresian.
Etymology. The epithet is in reference to the similarity of
this species to Hystrichokolpoma? incertum Michoux 1985.
Diagnosis. A species of Dapsilidinium in which some,
but not all, of the processes are significantly broader
than others, though the position of the broader process-
es appears not to be consistent.
Size. Central body maximum diameter 30‒32 μm; length
of processes up to about 15 μm; two specimens measured.
Age. LO: late Lutetian.
Remarks. This species is distinctive in having a more or less
bimodal variation in process widths, with a few being
distinctly broader than the others. The position and
number of broad processes vary from specimen to speci-
men. One or more of the broader processes on each spe -
cimen may be bifurcate distally down to about mid-
length. In Hystrichokolpoma? incertum, the larger processes
are consistently the precingulars and the antapical. In other
species of Dapsilidinium, such as Dap silidinium pseudocol-
ligerum and Dapsilidinium simplex, the processes are more
or less uniformly developed. In species of Diphyes, only the
antapical process is relatively large.
Dapsilidinium simplex (White 1842) Bujak et al. 1980
(Plate 5, fig. 14)
1842 Xanthidium tubiferum var. simplex White,
p. 38, plate 4, fig. 10.
1946 Hystrichosphaeridium simplex (White) –
Deflandre, card 934.
1969 Polysphaeridium? simplex (White) – Davey &
Williams, p. 7.
1980 Dapsilidinium simplex (White) – Bujak et al., p. 28.
Age. LO: Bartonian. Not plotted.
Genus Deflandrea Eisenack 1938
Type. Eisenack 1938, text-fig. 6, as Deflandrea phos-
phoritica.
1938 Deflandrea Eisenack, p. 187.
Remarks. Deflandrea is a peridiniacean cyst that is charac-
terised by its latideltaform intercalary (I2a) archaeopyle.
The broad shape is a consistently striking feature of the
periarchaeo pyle, but not always of the endoarchaeopyle.
Deflandrea borealis sp. nov.
(Plate 5, figs 17–20)
Holotype. Plate 5, fig. 19 from a cuttings sample at
3095‒3105 m in Rut H-11, GSC type collection no.
137964, sample P39397, slide 01, co-ordinates 18.9 ×
98.6, England Finder T41/2. Length (including horns)
57 μm, width 55 μm. The age determined for the
sample from which the holotype was recovered is Tha -
netian (Late Paleocene).
Etymology. The epithet is from the Latin borealis (north-
ern) in reference to the northern occurrence of at least
the type material of this species.
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41
Diagnosis. A relatively small, squat and generally rounded
species of Deflandrea with a scabrate to granulate wall and
a latiform archaeopyle, the operculum of which com -
monly remains attached posteriorly.
Size. Length (including horns) 46‒59 μm, width 46‒55
μm; three specimens measured.
Age. LO: latest Priabonian.
Deflandrea denticulata Alberti 1959
(Plate 5, figs 15, 16)
1959 Deflandrea denticulata Alberti, p. 102, 103,
text-fig. 1.
Age. LO: Ypresian. Not plotted.
Remarks. Deflandrea denticulata (Alberti 1959, text-fig.
1) is unusual for a species of this genus in having a peri-
cyst with long slender apical and antapical horns. The
periphragm is covered with small, slender spines. Al -
though the archaeopyle appears to be latideltaform, it is
impossible to be definite without examining the holo-
type. Alberti (1959) recorded Deflandrea denticulata
from Lower Eocene sediments of Volgograd, Russia,
the Oebisfelde Borehole in Germany, and possibly
from Belgium. His observations fit the predicted LO in
this study for a taxon with a morphology intermediate
between Cerodinium and Deflandrea.
Deflandrea galeata (Lejeune-Carpentier 1942)
Lentin & Williams 1973
(Plate 6, fig. 1)
1942 Peridinium galeata Lejeune-Carpentier,
p. B186–B188, figs 15–20.
1973 Deflandrea galeata (Lejeune-Carpentier) –
Lentin & Williams, p. 41.
Age. LO: Maastrichtian.
Remarks. The line-drawing of the dorsal view of the
holotype of Deflandrea galeata (Lejeune-Carpentier
1942, fig. 15) clearly shows a latideltaform hexa 2a
archae o pyle. This interpretation is confirmed in Le -
jeune-Carpentier & Sarjeant (1981, p. 18, 19, who
refer to the archaeopyle as “single-plate intercalary (type
I/I) of broad-hexa type and formed by the loss of para-
plate 2a”. In every other respect, the species has the
typical morphology of Cerodinium. It is unusual to find
forms with a latideltaform 2a in rocks of this age.
Deflandrea majae (Schiøler 1993) comb. nov.
(Plate 6, fig. 2)
1993 Isabelidinium majae Schiøler, p. 108, 110,
plate 1, figs 1–6; text-fig. 4a, b.
Age. LO: latest Maastrichtian.
Remarks. Schiøler (1993) noted that this species has a
latideltaform archaeopyle with a transverse archaeopyle
index (TAI) considerably higher than 0.5. Schiøler
(1993, p. 110) stated that although these characters are
“… typical of the genus Deflandrea … as the new
species lacks any signs of a paracingulum, referral to the
latter genus is precluded.” In our view, archaeopyle
shape is critical in diagnosing Deflandrea and similar
genera and that the presence or absence of a cingulum
is not significant. Therefore, this species is transferred
herein to Deflandrea. However, M. Pearce (personal
communication 2015) has pointed out that the archaeo -
pyles of specimens otherwise attributable to this species
show a wide variation in archaeopyle shape. For exam-
ple he has observed specimens of Deflan drea majae with
stenodeltaform archaeopyles; we would recommend that
such forms be included in another genus. As noted under
Alterbidinium, a de tailed re-evaluation of archaeo pyle
shapes in Deflan drea and similar genera in relation to
the taxonomy of the group is clearly needed but beyond
the scope of the present work.
Deflandrea oebisfeldensis Alberti 1959
(Plate 6, fig. 3)
1959 Deflandrea oebisfeldensis Alberti, p. 95, 96,
plate 8, figs 10–13.
Age. LO: early Ypresian.
Remarks. The antapical margin of the pericyst in De -
flandrea oebisfeldensis is broad, with the left and right
antapical horns being at the intersection with the poste-
rior lateral sides and thus broadly separated.
Deflandrea phosphoritica Eisenack 1938
(Plate 6, fig. 4)
1938 Deflandrea phosphoritica Eisenack, p. 187,
text-fig. 6.
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42
1965b Deflandrea granulosa Cookson & Eisenack,
p. 122, plate 11, figs 8, 9.
1965 Deflandrea heterophlycta forma pusulosa Rozen,
p. 293, 294, plate 1, figs 3, 4; text-fig. 2.
1966 Deflandrea menendezii Pöthe de Baldis,
p. 223, plate 2, fig. a.
1973 Deflandrea heterophlycta subsp. pusulosa
(Rozen) – Lentin & Williams, p. 41.
Age. LO: latest Chattian.
Genus Dinogymnium Evitt et al. 1967
Type. Evitt et al. 1967, plate 1, figs 21–23, plate 2, fig.
5, text-figs 16–18, as Dinogymnium acuminatum.
1967 Dinogymnium Evitt et al., p. 4–8.
Remarks. Lentin & Vozzhennikova (1990) subdivided
the dinogymnioids into genera that are separated on
overall outline, relative size of the episome and hypo-
some, and surface ornamentation. As a consequence,
these authors transferred several species previously in -
cluded in Dinogymnium into the new genera. Fensome
et al. (2009, p. 27) provided a synopsis for Dino gym -
nium that is followed here.
Dinogymnium longicorne (Vozzhennikova 1967)
Harland 1973
(Plate 6, fig. 11)
1967 Gymnodinium longicorne Vozzhennikova,
p. 46, plate 1, fig. 8; plate 3, fig. 6; plate 4,
figs 6a, b, 7.
1967 Gymnodinium curvatum Vozzhennikova,
p. 43, plate 1, figs 10–12; plate 4, figs 2, 3.
1973 Dinogymnium longicorne (Vozzhennikova) –
Harland, p. 678.
Age. LO: early Campanian.
Remarks. Dinogymnium longicorne is elongate, with an
episome that is considerably longer than the hyposome.
According to Lentin & Vozzhennikova (1990, p. 19),
the length varies between 62 and 91 μm, the width
between 21 and 38 μm.
Genus Diphyes Cookson 1965 nom. cons.
Type. Deflandre & Cookson 1955, plate 7, fig. 3, as
Hystrichosphaeridium colligerum.
1965 Diphyes Cookson, p. 85; name illegitimate.
1970 Lingulasphaera Drugg, p. 817.
2000 Diphyes Cookson nom. prop. cons. Harris &
Fensome, p. 281, 282.
Remarks. The conservation proposal of the name Diphyes
Cookson (Harris & Fensome 2000) was ratified at the
2005 Botanical Congress. We agree with the synopsis of
Diphyes in Fensome et al. (2009, p. 28), with one excep-
tion regarding the processes. Some specimens show an
intratabular organisation, with about four processes on
each pre- and postcingular plate.
Diphyes brevispinum Bujak 1994
(Plate 6, figs 5, 6)
1994 Diphyes brevispinum Bujak, p. 121, 123, plate
2, figs 4–6.
Age. LO: Ypresian.
Remarks. Diphyes brevispinum has short conical to subcon-
ical, rarely tapering processes and an inflated antapical
process that is similar to that of Diphyes ficusoides.
Diphyes colligerum (Deflandre & Cookson 1955)
Cookson 1965
(Plate 6, fig. 12)
1955 Hystrichosphaeridium colligerum Deflandre &
Cookson, p. 278, 279, plate 7, fig. 3.
1965 Baltisphaeridium colligerum (Deflandre &
Cookson) – Downie & Sarjeant, p. 88.
1994 Diphyes pseudoficusoides Bujak, p. 123, 125,
plate 2, figs 2, 3.
Age. LO: Lutetian.
Remarks. Following Fensome et al. (2009, p. 30), Di -
phyes pseudocolligerum should be considered a junior
synonym of Diphyes colligerum because the size of the
antapical process is similar in both holotypes.
Diphyes ficusoides Islam 1983a
(Plate 6, figs 7, 8)
1983a Diphyes ficusoides Islam, p. 338, plate 2, figs 8, 9.
Age. LO: middle Lutetian.
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Genus Disphaerogena Wetzel 1933a
Type. Wetzel 1933a, plate 4, fig. 34, as Disphaerogena
carposphaeropsis.
1933a Disphaerogena Wetzel, p. 51.
1976 Cyclapophysis Benson, p. 192.
1981 Plethysyrinx Sarjeant, p. 106.
Remarks. The synopsis of Disphaerogena provided in
Fensome et al. (2009, p. 30) is accepted, including that
Cy clapophysis is a taxonomic junior synonym of the genus.
Disphaerogena carposphaeropsis Wetzel 1933a
(Plate 6, fig. 9)
1933a Disphaerogena carposphaeropsis Wetzel, p. 51,
plate 4, fig. 34.
1976 Cyclapophysis monmouthensis Benson, p. 183,
plate 1, figs 9–12; plate 2, fig. 1.
Age. LO: latest Maastrichtian.
Remarks. When Sarjeant (1985a, p. 141, 142) emend-
ed the diagnosis of Disphaerogena carposphaeropsis, he
considered Cyclapophysis monmouthensis to be a taxo-
nomic junior synonym of the species.
Genus Eatonicysta Stover & Evitt 1978
Type. Morgenroth 1966a, plate 3, fig. 11, as Canno -
sphae ropsis ursulae.
1978 Eatonicysta Stover & Evitt, p. 41.
Synopsis. Gonyaulacacean (leptodinioid) cysts that are
chorate, with a spheroidal central body. Holocavate.
Central body bearing 17 to 23 hollow or solid, fibroid
mesotabular processes, which are distally connected by a
fenestrate to reticulate to irregular open-mesh, net-like
ectophragm. Archaeopyle apical, with formula A(1–4´),
operculum free.
Description. The holocavate cyst has an autophragm
bearing mesotabular processes with expanded distal
extremities that merge to form a perforate membranous
to reticulate to trabeculate ectophragm. The size of the
mesh or perforations shows considerable variation. A
leptodinioid tabulation of 3–4´, 6´´´´, 0–6c, 5´´´,
1´´´´, 0s is indicated by the processes. Based on process
size, plate 1´´ would be wider than 6´´.
Remarks. Eatonicysta is characterised by a membranous
ectophragm that may be reticulate or broken down to
form a trabeculate network.
Eatonicysta furensis (Heilmann-Clausen in Heil mann-
Clausen & Costa 1989) Stover & Williams 1995
(Plate 6, fig. 13)
1989 Eatonicysta ursulae subsp. furensis Heilmann-
Clausen in Heilmann-Clausen & Costa, p.
466, plate 11, figs 3, 5, 7.
1995 Eatonicysta furensis (Heilmann-Clausen in
Heilmann-Clausen & Costa) – Stover &
Williams, p. 104.
Age. LO: late Ypresian.
Remarks. Eatonicysta furensis has much shorter, usually
broader, funnel-shaped processes than Eatonicysta ursulae.
Also, the processes grade imperceptibly into the ecto -
phragm, which is divided into areas reflecting the indivi -
dual plates, rather than forming a continuous network.
Eatonicysta ursulae (Morgenroth 1966a) Stover &
Evitt 1978
(Plate 6, fig. 14)
1966a Eatonicysta ursulae Morgenroth, p. 20, plate
3, figs 11, 12.
1966a Membranilarnacia reticulata Williams & Downie,
p. 220, 221, plate 24, figs 4, 6; text-fig. 59.
1967 Membranilarnacia dictyophora Agelopoulos,
p. 49, 50, plate 12, figs 3, 4, 6.
1969 Membranilarnacia ursulae (Morgenroth) – de
Coninck, p. 43.
1978 Eatonicysta ursulae (Morgenroth) – Stover &
Evitt, p. 41.
Age. LO: earliest Lutetian.
Remarks. Williams & Downie (1966a) recorded two
variants of Eatonicysta ursulae (as Membranilarnacia re -
ticulata, a synonym of Eatonicysta ursulae) from the
Ypresian London Clay from southern England. One
lacked cingular processes, the other had four cingular
processes. Whether or not this variation is stratigraphi-
cally significant remains to be confirmed.
Genus Enneadocysta Stover & Williams 1995
Type. Gerlach 1961, plate 28, fig. 14, as Baltisphae r -
idium pectiniforme.
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44
1994 Enneadocysta Stover & Williams in Bujak,
p. 119; name not validly published.
1995 Enneadocysta Stover & Williams, p. 108, 109.
2007 Enneadocysta Stover & Williams – emend.
Fensome et al., p. 394.
Remarks. We follow the emendation of Fensome et al.
(2007, p. 394) in our concept for this genus and agree
that it is areoligeracean.
Enneadocysta magna Fensome et al. 2007
(Plate 7, figs 1, 2)
2007 Enneadocysta magna Fensome et al., p. 394,
396, plate 1, figs 1–20; plate 2, figs 1–19;
text-figs 5A, B, 6A–E.
Age. LO: latest Rupelian.
Remarks. Labrador Margin specimens of Enneadocysta
magna can have process clusters on individual plates.
Genus Eocladopyxis Morgenroth 1966a
Type. Morgenroth 1966a, plate 3, figs 2, 3, as Eoclado -
pyxis peniculata.
1966a Eocladopyxis Morgenroth, p. 7.
Remarks. The synopsis provided by Fensome et al.
(2009, p. 31) takes into account that Eocladopyxis is a
goniodomacean (pyrodinioid) cyst and identifies all the
plates involved in the formation of the archaeopyle.
Eocladopyxis peniculata Morgenroth 1966a
(Plate 7, fig. 3)
1966a Eocladopyxis peniculata Morgenroth, p. 7, 8,
plate 3, figs 2, 3.
Age. LO: late Ypresian.
Genus Evittosphaerula Manum 1979 emend.
Damassa 1997
Type. Manum 1979, plate 2, figs 3, 4, as Evittosphae -
rula paratabulata.
1979 Evittosphaerula Manum, p. 242, 243.
1997 Evittosphaerula Manum – emend. Damassa,
p. 161–163.
Evittosphaerula? foraminosa sp. nov.
(Plate 6, figs 15–20)
Holotype. Plate 6, figs 19, 20 from a cuttings sample at
2130–2140 m in North Leif I-05, GSC type collection
no. 138159, sample YD17600, slide 03, co-ordinates
44.9 × 15.5, England Finder M43/0. Pericyst length
82.5 μm, width 90 μm. The age determined for the
sample from which the holotype was recovered is
Ypresian.
Etymology. From the Latin foraminosus, meaning ‘full of
holes’.
Description. A species of gonyaulacalean cysts in which
only broad strips of membrane representing the sutures
are preserved. The tabulation appears to be gonio doma -
ceans, with a five-sided antapical plate reflecting a quin-
queform hypocystal tabulation. At the apex, sutural strips
come together to form a short apical horn.
Size. Diameter 68–90 μm; four specimens in dorso-
ventral orientation measured, but excluding the holo-
type as it is oriented apically–antapically.
Age. LO: Ypresian.
Remarks. This species has a very distinctive structure,
represented by strips of membrane reflecting the sutu-
res only; the internal area of each plate is represented by
a hole. Thus the species is reminiscent of the late Oligo -
cene to early Miocene species Evittospaherula paratabu-
lata, but differs superficially in having an apical horn,
an apparently much narrower cingulum, and broader
sutural membranes. More fundamentally, one speci-
men (Plate 6, fig. 18) appears to have a five-sided ant -
apical plate, suggesting a goniodomacean affinity (Fen -
some et al. 1993). The tabulation described by Manum
(1979) for Evittosphaerula paratabulata, the type of the
genus, is clearly gonyaulacacean. As this new species is
strikingly similar, albeit perhaps superficially, to Manum’s
species, and there is not sufficient material to describe
its tabulation in full, it is questionably assigned to
Evittosphaerula. The new species is also strikingly simi-
lar to both Hapsocysta susanae, described by Duxbury
(2002) from the Albian of the central North Sea, and
Chaenos phaerula magnifica, described by Damassa (1997)
from the late Oligocene of the Norwegian Sea. However,
the tabulation of both those species is clearly sexiform,
and neither has horns.
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45
Genus Fibrocysta Stover & Evitt 1978
Type. Cookson & Eisenack 1965b, plate 16, fig. 8, as
Cordosphaeridium bipolare.
1978 Fibrocysta Stover & Evitt, p. 155.
Synopsis. Gonyaulacacean (cribroperidinioid) cysts that
are chorate, with a longitudinally elongate ovoidal
central body, with a protrusion at the apical and antapi-
cal poles. Acavate or cornucavate, with a fibrous wall.
Processes numerous, nontabular to indistinctly tabular;
they are hollow and fibrous or solid and generally of
uniform size. Archaeopyle precingular, with formula
P3´´, operculum free.
Remarks. In wall structure and archaeopyle type, Fibro -
cysta is very similar to Turbiosphaera. However, process-
es in Tur bio sphaera are wider, especially apically and
antapically, and in Turbiosphaera, the cingulum is
marked by a membrane. The remarkable similarities
be tween the two genera suggest that Fibrocysta, like
Turbiosphaera, has a cribroperidinioid tabulation.
Fibrocysta bipolaris (Cookson & Eisenack 1965a)
Stover & Evitt 1978
(Plate 7, fig. 4)
1965a Cordosphaeridium bipolare Cookson &
Eisenack, p. 135, plate 16, figs 7, 8.
1969 Lanternosphaeridium bipolare (Cookson &
Eisenack) – de Coninck, p. 38.
1969c Amphorosphaeridium bipolare (Cookson &
Eisenack) – Davey, p. 35.
1978 Fibrocysta bipolaris (Cookson & Eisenack) –
Stover & Evitt, p. 155.
Age. Local acme early Ypresian.
Remarks. The cingulum of the holotype of Fibrocysta
bipo laris (Cookson & Eisenack 1965a, plate 16, fig. 8)
appears to be delineated by a single row of processes.
Genus Gillinia Cookson & Eisenack 1960a
Type. Cookson & Eisenack 1960a, plate 3, fig. 4, as
Gillinia hymenophora.
1960a Gillinia Cookson & Eisenack, p. 11, 12.
Synopsis. Small, proximate, slightly elongate cysts with
two membranous wings, one on each side of the apical
archaeopyle. Tabulation partially delineated by ridges,
which commonly define the cingulum and a sulcal area
that is considerably broader posteriorly.
Remarks. Following Fensome et al. (1993, p. 73), Gil -
linia probably has a cladopyxiacean tabulation.
Gillinia hymenophora Cookson & Eisenack 1960a
(Plate 7, fig. 12)
1960a Gillinia hymenophora Cookson & Eisenack, p.
12, plate 3, figs 4–6; text-fig. 5.
Age. LO: late Campanian.
Genus Ginginodinium Cookson & Eisenack 1960a
Type. Cookson & Eisenack 1960a, plate 2, fig. 9, as
Gin gi no dinium spinulosum.
1960a Ginginodinium Cookson & Eisenack, p. 7.
Remarks. In their emendation of Ginginodinium, Len tin
& Williams (1976, p. 95, 96) noted that the archaeopyle
is compound, involving the three anterior intercalary
plates and three of the precingular plates (3´´–5´´).
However, the archaeopyle may be formed from loss of
only the three anterior intercalary plates. There may also
be a series of successive stages, until all three intercalaries
are lost and three of the precingulars remain attached to
the main cyst solely along the cingular margin. This
variability in archaeopyle type often makes assignment
of species to the genus difficult.
Ginginodinium? flexidentatum sp. nov.
(Plate 7, figs 5–11)
Holotype. Plate 7, fig. 11, from a cuttings sample at
1815–1825 m in Bjarni O-82, GSC type collection no.
138070, sample P39715, slide 01, co-ordinates 3.6 ×
102.8, England Finder C32/4. Pericyst length 84 μm,
width 73 μm, endocyst length 61 μm, width 67 μm,
length of processes up to 2.5 μm. The age determined
for the sample from which the holotype was recovered
is early Ypresian.
Etymology. The epithet is from the Latin flexibilis, mean-
ing bendable and dentatus, meaning toothed or pointed.
Diagnosis. A species of Ginginodinium with one or more
lateral horns or bulges, a flexible folded wall, and no
clearly demarked cingulum. The ornament is variable
but typically denticulate.
Bulletin36.qxp_Bulletin 36 19/12/16 13.40 Side 45
46
Description. Pericyst outline pentagonal, with a well-
developed apical and two antapical horns, one of which
is slightly shorter than the other, and two lateral protu-
berances. Generally cornucavate but can, in part or
whole, be circumcavate. The endocyst is pentagonal.
Pericyst ornamentation varies from finely perforate to
verrucate to bearing processes. The archaeopyle in -
volves the loss of the 2a plate, which remains attached
to the 4´´ plate, and sometimes the partial detachment
of the 1a and 3a plates. One margin of the cingulum is
commonly indicated by a fold.
Size. Pericyst length 61–87 μm (mean 73 μm), pericyst
width 56–73 μm (mean 63 μm), endocyst length 43–
61 μm (mean 51 μm), width 49– 67 μm (mean 53 μm),
process length up to about 2.5 μm.
Age. LO: late Ypresian.
Remarks. The shapes of the pericyst and endocyst are
relatively stable in Ginginodinium? flexidentatum, but
there are differences between specimens in degree of
cavation, which can range from cornucavate to narrow-
ly circumcavate. There are also variations in the shape
of the horns, especially the apical: this is generally
rounded distally but may be acuminate. The antapical
horns are acuminate to rounded distally; the lateral
horns are primarily just protuberances. Pericyst orna-
mentation is variable, sometimes even on the same
specimen. The periphragm can be perforated and orna-
mented with small verrucae and processes, the latter of
variable length. Processes are slender and distally bifid.
This species is included in Ginginodinium only provi-
sionally because of the uncertainty over the archaeopyle.
Genus Glaphyrocysta Stover & Evitt 1978
Type. Cookson 1965, plate 11, fig. 4, as Cyclonephe -
lium retiintextum.
1978 Glaphyrocysta Stover & Evitt, p. 49, 50.
Remarks. The generic concept of Glaphyrocysta, as ex -
pressed in the emendation by Fensome et al. (2009, p.
32), is followed here. The genus is sometimes abundant
in middle Eocene sections in Labrador Margin wells.
Glaphyrocysta divaricata (Williams & Downie
1966a) Stover & Evitt 1978
(Plate 7, figs 15, 16)
1966a Cyclonephelium divaricatum Williams & Dow -
nie, p. 223, 224, plate 25, fig. 1; text-fig. 60.
1978 Glaphyrocysta divaricata (Williams &
Downie) – Stover & Evitt, p. 50.
Age. LO: earliest Lutetian. See also Remarks.
Remarks. A species of Glaphyrocysta with numerous pro -
cesses variously interconnected along their length, but
with a preponderance of distal free ends. In Glaphyro -
cysta divaricata, unlike in Glaphyrocysta ordinata and Gla -
phyrocysta retiintexta, the processes do not form distinct
complexes. Although this taxon has a Lutetian LO, it is
most abundant in the Paleocene and is often very
common to dominant in assemblages from that epoch.
Glaphyrocysta exuberans (Deflandre & Cookson
1955 ex Eaton 1976) Stover & Evitt 1978
(Plate 7, fig. 14)
1955 Cyclonephelium exuberans Deflandre &
Cookson, p. 255; name not validly published.
1976 Cyclonephelium exuberans Deflandre &
Cookson ex Eaton, p. 255, 256.
1978 Glaphyrocysta exuberans (Deflandre &
Cookson ex Eaton) – Stover & Evitt, p. 50.
Age. LO: Priabonian.
Remarks. Glaphyrocysta exuberans has a well-developed,
often perforate, marginate ectophragm, supported by
solid, slender processes.
Glaphyrocysta retiintexta (Cookson 1965) Stover
& Evitt 1978
(Plate 7, fig. 17)
1965 Cyclonephelium retiintextum Cookson, p. 88,
plate 11, fig. 4.
1978 Glaphyrocysta retiintexta (Cookson) – Stover
& Evitt, p. 50.
Age. LO: Priabonian.
Remarks. Glaphyrocysta retiintexta shows an ambital devel-
opment of distal trabecula but has minimal membrane
development. The general absence of membranes and the
distal connections between plate complexes distinguish
this species from Glaphyrocysta intricata. Fensome et al.
(2009, p. 34) noted that the morphology of Glaphyrocysta
retiintexta was very similar to that shown in the drawing
Bulletin36.qxp_Bulletin 36 19/12/16 13.40 Side 46
47
of the holotype of Glaphyrocysta pastielsii (Deflandre &
Cookson 1955) Stover & Evitt 1978 by Pastiels (1948,
plate 5, fig. 15). But Fensome et al. (2009) also stated that
photographs of the holotype of Glaphyrocysta pastielsii ap -
peared to possess a morphology more like that of Gla -
phy rocysta divaricata. Accordingly they recommended
restricting the name Glaphyrocysta pastielsii to the holo-
type, a proposal followed here.
Glaphyrocysta texta (Bujak 1976) Stover & Evitt 1978
(Plate 8, fig. 1)
1976 Cyclonephelium texta Bujak, p. 110, plate 3,
figs 6–11; text-fig. 3G, H.
1978 Glaphyrocysta texta (Bujak) – Stover & Evitt,
p. 50.
Age. LO: Priabonian.
Remarks. Glaphyrocysta texta has distinctive process com -
plexes on most of the pre- and postcingular plates, and
apparently one broader complex on the single antapical
plate. The contabular complexes consist of slender in -
tratabular processes, distally united to form clypeate plat-
forms that have ragged margins. The 3´´ and 3´´´ plates
sometimes have a single process rather than a process
complex. Adjacent process complexes are connected distal-
ly by trabecula.
Glaphyrocysta vicina (Eaton 1976) Stover & Evitt
1978
(Plate 8, fig. 2)
1976 Cyclonephelium vicinum Eaton, p. 260, 261,
plate 8, figs 4, 5; text-fig. 13.
1978 Glaphyrocysta vicina (Eaton) – Stover & Evitt,
p. 50.
Age. LO: Lutetian.
Remarks. Glaphyrocysta vicina has a marginal, perforate
pericoel, with the periphragm remaining close to the
endophragm. Both features are unusual in species of
Gla phy rocysta. Species of Membranophoridium lack pro -
cesses supporting the periphragm.
Genus Habibacysta Head et al. 1989
Type. Head et al. 1989, plate 4, figs 1, 2, 5, 6, as Habi -
bacysta tectata.
1989 Habibacysta Head et al., p. 457, 458.
Synopsis. Proximate spheroidal gonyaulacean cysts. Ata -
bulate. Autophragm bearing short columns that distally
are sometimes united by an entire, perforate or reticulate
layer. Archaeopyle precingular, with formula P3´´, opercu-
lum free.
Remarks. Head (1994, text-fig. 3) showed how the
nature of the autophragm differentiates Habibacysta
from Bitectatodinium, Filisphaera and Tectatodinium.
All four genera are proximate, atabulate gonyaulacacean
cysts with precingular archaeopyles.
Habibacysta tectata Head et al. 1989
(Plate 7, fig. 13)
1989 Habibacysta tectata Head et al., p. 458, plate
4, figs 1–6, 9, 10.
Age. LO: earliest Zanclean.
Genus Hapsocysta Davey 1979
Type. Eisenack & Cookson 1960, plate 3, fig. 6, as
Can nosphaeropsis peridictya.
1979 Hapsocysta Davey, p. 556.
Synopsis. A chorate to camocavate gonyaulacacean (go ny -
au la coidean) cyst, with a subrounded to ovoidal central
body, which is surrounded by a periphragm that may be
trabeculate, forming an open network, or filled by a
per forate membrane. Tabulation indicated by the trabe -
cula, which on membranous taxa occur as ridges.
Archaeo pyle precingular, with formula P3´´, operculum
free.
Remarks. Fensome et al. (1993, p. 89) considered Hap -
socysta to have a cribroperidinioid tabulation. However,
Nøhr-Hansen (1993, p. 71, 72) showed that the taxon,
which he named Hapsocysta? benteae, had gonyaula-
coidean tabulation. Nøhr-Hansen (1993) provisionally
included the species in Hapsocysta, because of the pres-
ence of the thin-walled periphragm.
Hapsocysta? benteae Nøhr-Hansen 1993
(Plate 8, fig. 12)
1993 Hapsocysta? benteae Nøhr-Hansen, p. 71, 72,
plate 25, figs 11, 12; text-figs 10a, b, 11a, b.
Age. LO: Cenomanian.
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48
Genus Heteraulacacysta Drugg & Loeblich Jr.
1967
Type. Drugg & Loeblich Jr. 1967, plate 1, fig. 8a–c, as
Heteraulacacysta campanula.
1967 Heteraulacacysta Drugg & Loeblich Jr.,
p. 183.
Remarks. The synopsis for Heteraulacacysta by Fen some
et al. (2009, p. 35) is followed.
Heteraulacacysta porosa Bujak in Bujak et al. 1980
(Plate 8, fig. 16)
1980 Heteraulacacysta porosa Bujak in Bujak et al.,
p. 62, plate 15, figs 10–13; text-fig. 14B, C.
Age. LO: Priabonian.
Remarks. Considerable uncertainty has prevailed over
the distinction between Heteraulacacysta leptalea Eaton
1976 and Heteraulacacysta porosa. In his diagnosis for
Heteraulacacysta leptalea, Eaton (1976, p. 305) stated:
“Circular fenestrations frequently developed in the prox-
imal area of the cingular crests, along with fine elongate
fenestrations aligned at right-angles to the margin of the
cyst body.” Under ‘Remarks’, Eaton commented that
small perforations occurred in the periphragm, compara-
ble to those in the proximal area of the cingular crests,
and commonly gave a punctate appearance to the wall.
In the succeeding sentence, Eaton (1976, p. 305) differ-
entiated Heteraulacacysta leptalea from Heteraulacacysta
campanula on the basis of its “…frequently punctuate
rather than positively ornamented surface to the cyst
body, and in exhibiting circular and elongate fenestra-
tions in the cingular crests.”
Bujak in Bujak et al. (1980, p. 62) differentiated
Heteraulacacysta porosa “… from all other described
species of Heteraulacacysta by its perforate periphragm.”
The illustrations of the holotype and another specimen
of Heteraulacacysta leptalea in Eaton (1976, plate 21,
figs 1, 2) clearly show perforations. Consequently, the
only consistent difference appears to be the presence of
elongate perforations on the cingular crests in Hete rau -
lacacysta leptalea. We do not know, however, whether
this feature is stratigraphically significant.
Genus Heterosphaeridium Cookson & Eisenack
1968
Type. Cookson & Eisenack 1968, text-fig. 4H, as He -
tero sphae ridium conjunctum.
1968 Heterosphaeridium Cookson & Eisenack, p 115.
Remarks. In their synopsis for Heterosphaeridium, Fen -
some et al. (2009, p. 35) recognised that it is an areolig-
eracean cyst with a spheroidal to broadly ovoidal central
body. Yun Hyesu (1981, p. 45, 46) provided an emend -
ed diagnosis that broadened the circumscription mainly
to include forms with hollow, open processes as well as
solid processes. Some of the specimens of Heteros phae r -
idium difficile observed in the present study have pro -
cesses that are perforated or circular (annulate) process
complexes that can be connected at various locations
along their length.
Heterosphaeridium bellii Radmacher et al. 2014
(Plate 8, figs 3, 4)
2014 Heterosphaeridium bellii Radmacher et al.,
p. 31–33, plate 1, figs 1–9.
Age. LO: late Campanian.
Remarks. Heterosphaeridium bellii differs from Hetero -
sphaeridium heteracanthum in having dolabrate pro cesses
that are not branching. The processes in Het e ro s phae r -
idium bellii may be connected proximally but are never
branched along their length. Radmacher et al. (2014)
considered the LO of Heterosphaeridium bellii to be
late Campanian to early Maastrichtian in the south-
western Barents Sea.
Heterosphaeridium difficile (Manum & Cookson
1964) Ioannides 1986
(Plate 8, fig. 8)
1964 Hystrichosphaeridium difficile Manum &
Cookson, p. 12–14, plate 3, figs 1–3, 7.
1986 Heterosphaeridium difficile (Manum &
Cookson) – Ioannides, p. 24.
Age. LO: early Santonian.
Remarks. Labrador Margin specimens of Heteros phae r -
idium difficile show extreme variation in the nature of the
processes. Some are perforate along their length or arched
so that the process walls merge distally. Others, as noted
under ‘Remarks’ for the genus, have process complexes on
the pre- and postcingular plates that are connected by
circular proximal membranes, as in Systematophora. In the
latter specimens, the cingular plates have linear process
complexes. In some specimens, the apical plates com -
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49
monly show a characteristic X-shaped crest as observed by
M. Pearce (personal communication 2015) and also
appear to have annulate process complexes. The antapical
plate is marked by an annulate complex that is broader
than those on the pre- and postcingular plates. The wall
of the central body is microreticulate.
Genus Histiocysta Davey 1969a
Type. Davey 1969a, plate 1, fig. 5, text-fig. 14A, B, as
Histiocysta palla.
1969a Histiocysta Davey, p. 138.
Remarks. This genus is similar to Corrudinium in its
shape and ornamentation, but tends to be smaller and
has an apical archaeopyle.
Histiocysta palla Davey 1969a
(Plate 7, figs 18–20)
1969a Histiocysta palla Davey, p. 138–140, plate 1,
figs 5, 6; text-fig. 14A, B.
Age. LO: late Campanian.
Genus Homotryblium Davey & Williams 1966a
Type. Davey & Williams 1966a, plate 12, fig. 5, as Ho -
mo tryblium tenuispinosum.
1966a Homotryblium Davey & Williams, p. 100.
Remarks. The synopsis provided by Fensome et al.
(2009, p. 35, 36) covers all the key morphological
attributes of Homotryblium.
Homotryblium abbreviatum Eaton 1976
(Plate 8, figs 9, 10)
1976 Homotryblium abbreviatum Eaton, p. 267,
268, plate 10, figs 2–4.
Age. LO: late Ypresian. Not plotted.
Remarks. As in the type material, the Labrador Margin
specimens of Homotryblium abbreviatum have short and
wide processes, but the surface ornamentation of the
central body can be smooth as well as granulate. This
species is not as common as Homotryblium tenuis pi no -
sum in the samples studied here.
Homotryblium tenuispinosum Davey & Williams
1966a
(Plate 8, figs 5–7)
1966a Homotryblium tenuispinosum Davey &
Williams, p. 101, 102, plate 4, fig. 11; plate
12, figs 1, 5, 7; text-fig. 21.
1966a Homotryblium pallidum Davey & Williams,
p. 102, 103, plate 12, figs 4, 6; text-fig. 22.
Age. LO: Bartonian; peak: latest Ypresian.
Genus Hystrichokolpoma Klumpp 1953
Type. Klumpp 1953, plate 17, figs 3, 5a, as Hystri cho -
kolpoma cinctum.
1953 Hystrichokolpoma Klumpp, p. 388.
Remarks. Fensome et al. (2009, p. 36) provided a de -
tailed synopsis for Hystrichokolpoma, which stipulated
among other morphologic attributes, that the genus
must have cingular processes.
Hystrichokolpoma cinctum Klumpp 1953
(Plate 8, fig. 11)
1953 Hystrichokolpoma cinctum Klumpp, p. 389,
plate 17, figs 3, 4, 5a–d.
Age. LO: Burdigalean according to Williams et al.
(2004), but we consider that the age range of this species
needs to be better constrained. Not plotted.
Remarks. A species of Hystrichokolpoma in which the pre-
and postcingular processes fill plates and have small
tubular extensions. There are generally two processes per
cingular paraplate.
Hystrichokolpoma globulus Michoux 1985
(Plate 8, figs 13–15)
1985 Hystrichokolpoma globulus Michoux, p. 143,
plate 1, figs 1–4, 12; text-fig. 2A, B.
Age. LO: late Ypresian. Not plotted.
Remarks. Michoux (1985, p. 143) compared Hystricho -
kol poma globulus to Hystrichokolpoma cinctum Klumpp
1953, from which it differs by having a much shorter
antapical process relative to the length of the cyst and
pre- and postcingular processes that do not branch into
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50
tubules distally. Also, the cingular and sulcal processes
are conical and there are three to five on each cingular
plate. The Labrador Margin specimens of Hystricho -
kolpoma globulus differ from the type material in not
having conical cingular plates and in there being only
three or less on each plate.
Genus Hystrichosphaeridium Deflandre 1937
Type. Ehrenberg 1838, plate 1, fig. 16, as Xanthidium
tubiferum.
1937 Hystrichosphaeridium Deflandre, p. 68.
Remarks. We concur with the synopsis provided by
Fensome et al. (2009, p. 38) except to note that individ-
ual processes proximally do not cover most of the
underlying plate and that the cingular and sulcal
processes are slender. The apical processes also tend to
be slender, especially the first and fourth. Variation in
process dimensions in Hystrichosphaeridium does not
approach the strong variation that is distinctive of
Hystrichokolpoma.
Hystrichosphaeridium quadratum sp. nov.
(Plate 8, figs 17–19)
Holotype. Plate 8, figs 17, 18, from a cuttings sample at
2770–2780 m in Gilbert F-53, GSC type collection no.
137988, sample P39505, slide 01, co-ordinates 11.3 ×
99.2, England Finder L29/3. Diameter of central body
44 μm, length of processes up to about 37 μm. The age
determined for the sample from which the holotype
was recovered is early Ypresian.
Etymology. From the Latin quadratus (four-cornered),
in reference to the rectangular distal endings of the
processes in this species.
Diagnosis. A species of Hystrichosphaeridium in which
the distal process endings are slightly flared and poly -
gonal, generally rectangular and commonly perforate.
The process bases are mesotabular and circular.
Size. Width of central body 36‒48 μm, length of cen -
tral body (when operculum in place) 46‒63 μm; pro -
cesses up to 39 μm long. Six specimens measured.
Age. LO: Selandian. Not plotted.
Remarks. This species differs from Hystrichos phaeri -
dium tubiferum in having rectangular endings to its
processes. It also resembles Hystrichokolpoma proprium,
but the holotype of that species has rounded polygonal
process bases that largely fill the plate (see Fauconnier
& Masure 2004, plate 40, figs 1–3), as is typical of the
genus Hystrichokolpoma. Hystrichos phae ridium salpin-
gophorum differs in lacking perforations in the distal
endings of the processes.
Hystrichosphaeridium tubiferum (Ehrenberg 1838)
Deflandre 1937
(Plate 8, fig. 20)
1838 Xanthidium tubiferum Ehrenberg, plate 1, fig. 16.
1937 Hystrichosphaeridium tubiferum (Ehrenberg) –
Deflandre, p. 68.
Age. LO: Lutetian.
Genus Hystrichosphaeropsis Deflandre 1935
Type. Deflandre 1935, plate 8, fig. 11, as Hystri cho -
sphaeropsis ovum.
1935 Hystrichosphaeropsis Deflandre, p. 232.
1937 Hystrichosphaera subgenus Hystrichosphaeropsis
(Deflandre) Deflandre, p. 67.
Remarks. Eisenack (1963b, p. 118) retained Hystricho -
sphaeropsis at generic rank. Fensome et al. (2009, p. 38)
provided a synopsis that allows for the inclusion of
circumcavate forms in Hystrichosphaeropsis.
Hystrichosphaeropsis perforata Schiøler 1993
(Plate 9, fig. 4)
1993 Hystrichosphaeropsis perforata Schiøler, p. 106,
plate 2, figs 4–8; text-fig. 3.
Age. LO: late Maastrichtian.
Hystrichosphaeropsis quasicribrata (Wetzel 1961)
Gocht 1976
(Plate 9, fig. 12)
1961 Triblastula quasicribrata Wetzel, p. 340, plate
2, fig. 3.
1976 Hystrichosphaeropsis quasicribrata (Wetzel) –
Gocht, p. 322.
Age. LO: late Maastrichtian.
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51
Genus Hystrichostrogylon Agelopoulos 1964
Type. Agelopoulos 1967, text-figs 1, 2, as Hystri cho stro -
gylon membraniphorum.
1964 Hystrichostrogylon Agelopoulos, p. 673, 674.
Hystrichostrogylon digitus sp. nov.
(Plate 9, figs 1–3)
Holotype. Plate 9, fig. 3, from a cuttings sample at
2435‒2445 m in Rut H-11, GSC type collection no.
137952, sample P39375, slide 01, co-ordinates 15.8 ×
105.0, England Finder Q48/1. Length of central body
(excluding cavation) 45 μm, width of central body
(excluding cavation) 38 μm; overall length of cyst 73
μm, overall width of cyst 72 μm. The age determined
for the sample from which the holotype was recovered
is Lutetian Priabonian.
Etymology. From the Latin digitus (finger), in reference
to the long extensions at the process endings. The
epithet is a noun in apposition.
Diagnosis. A species of Hystrichostrogylon in which the
bi- and trifurcations of the processes constitute long,
fine, cylindrical distal branches generally one-third to
one half the length of the process stem. The distal
branches tend to be perpendicular to the process stem.
Size. Length of central body (excluding cavation)
40‒45 μm, width of central body (excluding cavation)
33‒40 μm; overall length of cyst 73‒82 μm, overall
width of cyst 66‒79 μm; 3 specimens measured.
Age. LO: Bartonian‒Priabonian. Not plotted.
Remarks. This species differs from other species of Hy -
s tric ho strogylon in having remarkably long, fine branch-
es at the ends of the processes.
Genus Impagidinium Stover & Evitt 1978
Type. Cookson & Eisenack 1965b, plate 12, figs 5–6,
as Leptodinium dispertitum.
1978 Impagidinium Stover & Evitt, p. 165, 166.
Synopsis. Proximate gonyaulacaceans with the S-type
ven tral organisation; in dorso-ventral view they are
subspheroidal to ellipsoidal. Tabulation clearly defined
by sutural septa or thickenings, although sometimes
these features are missing between plates bordering the
sulcus or the cingulum. Sometimes the septum or
thickening between the 3´ and 4´ plates is reduced or
absent. Cingulum and sulcus clearly defined. Archae -
opyle precingular, with formula P3´´, operculum free.
Remarks. The inclusion of Impagidinium in the sub -
family Gonyaulacoideae by Fensome et al. (1993, p.
92) reflects an appreciation of the commonly triangular
shape of the 6´´ plate. If a separate reflected plate, the
4´ is generally not in contact with the anterior sulcal
plate.
Impagidinium victorianum (Cookson & Eisenack
1965b) Stover & Evitt 1978
(Plate 9, figs 9–11, 13, 14)
1965b Leptodinium victorianum Cookson &
Eisenack, p. 123, plate 12, figs 8, 9.
1978 Impagidinium victorianum (Cookson &
Eisenack) – Stover & Evitt, p. 166.
Age. LO: intra late Maastrichtian.
Remarks. Cookson & Eisenack (1965b, p. 123) noted
that Impagidinium victorianum described from the Late
Eocene differs from Impagidinium dispertitum in its
“larger size, spherical form, and the constant absence of
the transverse dividing between the upper two plates of
the ventral field….” Where the anterior ventral side of
the specimens in this study can be observed, Impag -
idinium victorianum also lacks the suture between what
are purportedly the upper two plates of the sulcus.
However, Labrador Sea specimens of Impagidinium
victorianum are generally smaller than the type materi-
al, which ranges from 80 to 120 μm in length and 80 to
123 μm in width. In contrast, the Labrador Sea speci-
mens vary from 60 to 71 μm in length and 56 to 80 μm
in width. Distal and proximal ends of the cingulum are
offset by approximately a cingulum width. The sutural
features may be ridges or septa but are consistent within
an individual specimen. Maximum height of the sutur-
al ridges or septa is 5 μm. In some specimens the septa
are perforated, the perforations aligned in single rows.
Whether those forms with perforations should be in -
cluded in a new species is dependent upon finding more
specimens.
An intriguing aspect of the occurrence of Im -
pagidinium victorianum in the Labrador Sea is that its
LO is consistently within the Maastrichtian. This may
reflect changing environments from deep-water to shal-
lower conditions, but this does not explain the absence
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52
of the species in deeper water parts of the Palaeogene.
One explanation advanced by M. Pearce (personal
com munication 2015) is that Impagidinium victori-
anum is a warmer-water species that migrated south
during cooler climatic conditions. If so, it is odd that it
did not migrate back during Paleocene warming, unless
opening of the Labrador–Baffin Seaway had already
generated a proto-Labrador Current.
Genus Impletosphaeridium Morgenroth 1966a
Type. Morgenroth 1966a, plate 10, fig. 5, as Impleto -
sphaeridium transfodum.
1966a Impletosphaeridium Morgenroth, p. 32.
1971 Ciliosphaeridium Grigorovich, p. 94.
1984c Laticavodinium Wilson & Sarjeant in
Sarjeant, p. 127.
Remarks. In their remarks on Impletosphaeridium, Fen -
some et al. (2009, p. 38) agreed with Islam (1993, p.
84, 85) that the archaeopyle of the paratype of Imple -
tosphaeridium transfodum, the ‘type species’, is probably
apical. But the morphology of the holotype remains
uncertain. To quote Fensome et al. (2009, p. 38): “This
genus is useful, if of dubious status, since it serves as a
repository for chorate forms whose general morphology
accords with a gonyaulacacean dinoflagellate affinity,
but whose archaeopyle is uncertain.”
Impletosphaeridium apodastum sp. nov.
(Plate 9, figs 5–8)
Holotype. Plate 9, fig. 5, from a cuttings sample at 4565–
4575 m in Hekja O-71, GSC type collection no.
137903, sample P18737, slide 01, co-ordinates 13.3 ×
108.0, England Finder N37/3. Central body length 30
μm, width 26 μm; length of processes up to 17 μm,
width less than 1 μm. The age determined for the sample
from which the holotype was recovered is late Danian.
Etymology. The epithet is from the Greek apodastos,
meaning separated or apportioned, in reference to the
bifurcations of the processes distally.
Diagnosis. A species of Impletosphaeridium with solid
delicate processes that divide distally into two relatively
long branches. The branches are bifurcate at the tips
and usually curve back towards the central body.
Description. This species has a rounded, small central
body that is granulate. Distally, the solid delicate pro -
cesses divide into two, relatively long branches, each of
which is bifurcate at its tip. These distal branches usual-
ly curve back towards the central body. The initial
bifurcations are generally one-quarter to one-third as
long as the main stem of the process but can be as much
as a half or as little as a quarter the length. There are
about 50 processes per specimen.
Size. Central body length 20–33 μm, width 18–27 μm,
length of processes 10–18 μm; six specimens measured.
Age. LO: Selandian.
Remarks. Impletosphaeridium apodastum is character -
ised by the high number of processes and their distinc-
tive distal extremities, which branch. Tips of the two
branch es are usually bifid. Occasional processes have
three branches. Because of the unknown nature of the
archaeo py le, this species is included in Impletosphaeridium.
Genus Isabelidinium Lentin & Williams 1977b
Type. Cookson & Eisenack 1958, plate 4, fig. 10, as
Deflandrea korojonensis.
1976 Isabelia Lentin & Williams, p. 56 (name ille-
gitimate).
1977b Isabelidinium Lentin & Williams, p. 167.
2009 Isabelidinium Lentin & Williams – emend.
Fensome et al., p. 39.
Remarks. In recent decades, there has been much de -
bate about the definition of Isabelidinium and similar
genera such as Manumiella and Chatangiella, all of
which have an isodeltaform, isothetaform and/or iso-
omegaform 2a plate whose partial or complete detach-
ment forms the archaeopyle. Species now included in
Chatangiella and Isabelidinium can have an isodelata -
form, isothetaform or iso-omegaform 2a plate, but
Manu miella almost always has an isodeltaform 2a plate.
Much confusion remains regarding generic circum-
scriptions, but a resolution as to how these should be
dealt with is beyond the scope of the present study. In
our view, generic distinctions should be based mainly
on variations in the shape and dimensions of the hexa
2a plate and secondarily on other expressions of tabula-
tion such as the cingulum and on the ornamentation.
Chatangiella is clearly distinguished from Isabelidi -
nium by its partite cingulum. Fensome et al. (2009)
emended Manumiella, restricting it to forms with one
symmetrically disposed antapical prominence. How -
ever, the antapex of the holotype of the type of the genus,
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53
Manumiella (originally Broomea) seelandica (Lange
1969, plate 3, fig. 3), is partly obscured so that its
precise morphology cannot be determined; and the
paratype (Lange 1969, plate 2, fig. 10) clearly has two
protuberances. Thorn et al. (2009) emended the diag-
nosis of Manumiella to allow for inclusion of a mes -
ophragm, which we would consider a non-diagnostic
feature at generic rank. However, these authors did in -
dicate that Manumiella was distinctive in being cir -
cumcavate, with Isabelidinium being bicavate. For further
comparisons, see under Alterbidinium.
Isabelidinium cooksoniae (Alberti 1959) Lentin &
Williams 1977b
1959 Deflandrea cooksoniae Alberti, p. 97, plate 9, figs 1–6.
1961b Isabelidinium belfastense Cookson & Eisenack,
p. 71, plate 11, figs 4–6.
1967 Australiella cooksoniae (Alberti) –
Vozzhennikova, p. 132.
1976 Isabelia cooksoniae (Alberti) – Lentin &
Williams, p. 57.
1976 Isabelia belfastensis (Cookson & Eisenack) –
Lentin & Williams, p. 57.
1977b Isabelidinium cooksoniae (Alberti) – Lentin &
Williams, p. 167.
1977b Isabelidinium belfastense (Cookson &
Eisenack) – Lentin & Williams, p. 167.
1992 Isabelidinium bujakii Marheinecke, p. 86, 87,
plate 18, figs 1–3; text-fig. 16.
Age. LO: Maastrichtian.
Remarks. See Fensome et al. (2009, p. 39) for a discus-
sion of this species.
Isabelidinium cretaceum (Cookson 1956) Lentin &
Williams 1977b
(Plate 9, figs 15, 16)
1956 Deflandrea cretacea Cookson, p. 184, 185,
plate 1, figs 1–4 (only).
1976 Isabelia cretacea (Cookson) – Lentin &
Williams, p. 57.
1977b Isabelidinium cretaceum (Cookson) – Lentin
& Williams, p. 167.
1983 Manumiella? cretacea (Cookson) – Bujak &
Davies, p. 161.
Age. LO: Maastrichtian.
Remarks. The holotype of this species (Cookson 1956,
plate 1, fig. 1 and Helby et al. 1987, fig. 42L) and some
of the other specimens illustrated in the protologue
(Cookson (1956, plate 1, figs 2–4) are all bicavate, with
an endocyst that is broader than long. This bicavation,
together with a general rounding of the apical and
antapical regions, are considered here to be characteris-
tic for Isabelidinium cretaceum. However, the presence
of a very short horn on two specimens (Cookson 1956,
plate 1, figs 3, 4), neither of which is the holotype, does
not exclude their retention in this species. This species
is retained in Isabelidinium because it is bicavate (see
discussion above under Isabelidinium).
Isabelidinium microarmum (McIntyre 1975)
Lentin & Williams 1977b
(Plate 9, fig. 17)
1975 Deflandrea microarma McIntyre, p. 65, plate
l, figs 5–8.
1976 Isabelia microarma (McIntyre) – Lentin &
Williams, p. 58.
1977b Isabelidinium microarmum (McIntyre) –
Lentin & Williams, p. 168.
Age. LO: early Campanian.
Remarks. The holotype of Isabelidinium microarmum
(McIntyre 1975, plate 1, figs 5, 6) has a deltaform
archae opyle as do all the other specimens (McIntyre
1975, plate 1, figs 6–8).
Genus Kiokansium Stover & Evitt 1978
Type. Tasch et al. 1964, plate 3, fig. 8, as Hystricho -
sphaeridium unituberculatum.
1978 Kiokansium Stover & Evitt, p. 167.
1979 Bacchidinium Davey, p. 555.
Remarks. Characteristic features of Kiokansium are the
precingular archaeopyle, which is formed from the loss
of plates 3´´ and 4´´, and the dirigate to cauliflorate
distal terminations of the numerous solid processes.
Kiokansium williamsii Singh 1983
1983 Kiokansium williamsii Singh, p. 150, plate 54,
figs 3–6.
Age. LO: Cenomanian.
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Genus Kleithriasphaeridium Davey 1974
Type. Davey 1974, plate 5, figs 1, 2, text-fig. 3, as Klei -
thriasphaeridium corrugatum.
1974 Kleithriasphaeridium Davey, p. 55, 56.
1976 Diversispinosa Benson, p. 184.
Age. LO: Coniacian.
Remarks. Fensome et al. (2009, p. 40) emended the diag-
nosis of Kleithriasphaeridium to include forms with
mesotabular, tubular processes that are open or closed
distally and “to circumscribe forms with a combination
precingular–apical archaeopyle, a type of opening that
occurs as an intraspecific variant of some species.”
These authors also stressed that Kleithriasphaeridium
does not have fibrous processes, thus differentiating it
from Cordosphaeridium.
Kleithriasphaeridium mantellii (Davey & Williams
1966a) comb. nov.
(Plate 9, fig. 18)
1966a Hystrichosphaeridium mantellii Davey &
Williams, p. 66, plate 6, fig. 6.
1973 Florentinia mantellii (Davey & Williams) –
Davey & Verdier, p. 191.
Age. LO: Coniacian.
Remarks. Davey & Williams (1966a, p. 66) stated that
“The periphragm of processes [is] slightly fibrous.” On
the same page they further noted that “… the peri -
phragm of the central body appears to be fairly heavily
granular but on closer examination the granules appar-
ently result from a fine reticulation” and that “An apical
archaeopyle appears always to be present.” Davey &
Verdier (1973, plate 4, figs 1, 3) re-illustrated the holo-
type of Kleithrias phaeridium (as Florentinia) mantellii
and concluded that the archaeopyle is precingular, re -
sulting from the loss of the 3´´ plate. Although the
processes may be slightly fibrous, it is not obvious in the
photographs. Thus the characteristic feature of the
species appears to be the or namentation of the main
body. Fensome & Williams (2005, p. 48) noted that
Florentinia includes “forms with simple tubular processes
such as Florentinia aculeata and Florentinia cooksoniae, as
well as more ‘classic’ Florentinia types, such as Florentinia
laciniata and Florentinia ferox, with more complicated
processes.” They recommended that taxa with simple
tubular processes and precingular rather than combina-
tion archaeopyles, such as Florent inia cooksoniae, should
be transferred to Kleithrias phaeri dium. Thus the new
com bination Kleithriasphaeridium mantellii is proposed
here.
Genus Laciniadinium McIntyre 1975
Type. McIntyre 1975, plate 4, figs 12, 13, as Lacini a -
dinium orbiculatum.
1975 Laciniadinium McIntyre, p. 70.
1984 Sinocysta He Chengquan, p. 769, 773.
Synopsis. A proximate peridiniacean (palaeoperidin-
ioidean) cyst that is subspherical to biconical and com -
pressed dorso-laterally; when biconical, single horns are
developed at the apical and antapical poles. Acavate.
Autophragm smooth or ornamented with granules or
echinae. Cingulum indicated by low ridges. Archaeo -
pyle combination intercalary-precingular, with formula
IP(1–3a + 3–5´´), operculum simple, attached along the
anterior margin of the cingulum.
Remarks. Laciniadinium differs from Palaeoperidinium
in having one rather than two antapical horns, and an
archaeopyle involving intercalary and precingular plates
only.
Laciniadinium arcticum (Manum & Cookson
1964) Lentin & Williams 1980
(Plate 9, figs 19, 20)
1964 Diconodinium arcticum Manum & Cookson,
p. 18, 19, plate 6, figs 1–4.
1980 Laciniadinium arcticum (Manum &
Cookson) – Lentin & Williams, p. 41.
1986 Laciniadinium williamsii Ioannides, p. 28,
plate 10, figs 1–6; plate 11, fig. 5.
Age. LO: late Maastrichtian.
Remarks. In their description of Laciniadinium (as
Diconodinium) arcticum, Manum & Cookson (1964)
stated that the autophragm was ornamented with
minute granules up to 0.5 μm in diameter. The size
range of the species was: length 50–73 μm, breadth 32–
53 μm. Ioannides (1986) erected the species Lacinia di -
nium williamsii, which can have a smooth to finely
or namented autophragm; Laciniadinium williamsii va -
ries from 39 to 54 μm in length and 31 to 43 μm in
width. Unfortunately, Ioannides (1986) did not com -
pare Laciniadinium williamsii to Laciniadinium ar c ti -
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55
cum, although he did differentiate it from Lacinia -
dinium orbiculatum McIntyre 1975 and Laciniadinium
biconiculum McIntyre 1975. The only difference that
we can determine for separating Laciniadinium arcti -
cum and Laciniadinium williamsii is on the size, but this
overlaps as demonstrated above. Accordingly, we herein
consider Laciniadinium williamsii to be a taxonomic
junior synonym of Laciniadinium arcticum.
Genus Lentinia Bujak in Bujak et al. 1980
Type. Bujak et al. 1980, plate 18, figs 7–9, text-fig.
18A–F, as Lentinia serrata.
1980 Lentinia Bujak in Bujak et al., p. 69.
Remarks. Fensome et al. (2009, p. 42) provided a com -
prehensive synopsis for Lentinia that includes reference
to the large 2a isodeltaform archaeopyle, which domi-
nates the dorsal epicystal area of the pericyst.
Lentinia serrata Bujak in Bujak et al. 1980
(Plate 10, fig. 9)
1980 Lentinia serrata Bujak in Bujak et al., p. 71,
72, plate 18, figs 7–12; text-figs 18A–F, 19.
Age. LO: latest Priabonian.
Genus Licracysta Fensome et al. 2007
Type. Fensome et al. 2007, plate 4, figs 9–12, as Licra -
cysta corymbus.
2007 Licracysta Fensome et al., p. 400, 402.
Remarks. Licracysta is an areoligeracean cyst with non -
tabular to penitabular processes, many dolabrate to
moderately licrate, that are absent from the dorsoven-
tral region. In Glaphyrocysta, the processes are not
licrate or dolobrate distally. Also, processes tend to be
longer relative to the size of the central body than in
Licracysta. A synopsis for Licracysta is provided by Fen -
some et al. (2009, p. 42).
Licracysta corymbus Fensome et al. 2007
(Plate 10, fig. 16)
2007 Licracysta corymbus Fensome et al., p. 402,
404, 406, 408; plate 4, figs 5, 6, 9–20; plate
5, figs 1–8, 12, 16, 20.
Age. LO: Rupelian.
Licracysta? semicirculata (Morgenroth 1966b)
Fensome et al. 2007
1966b Cyclonephelium semicirculatum Morgenroth,
p. 9, 10, plate 2, figs 3, 4.
1978 Areoligera semicirculata (Morgenroth) – Stover
& Evitt, p. 18.
2007 Licracysta? semicirculata (Morgenroth) – Fen -
some et al., p. 408.
Age. LO: latest Rupelian.
Remarks. The distribution of the process complexes in
Licracysta? semicirculata is reminiscent of the Late
Cretaceous forms of Areoligera, in which the mid-
ventral surface lacks ornamentation, ventral processes
being restricted to ambital linear complexes.
Genus Lingulodinium Wall 1967
Type. Deflandre & Cookson 1955, plate 9, fig. 6, as
Hystrichosphaeridium machaerophorum.
1967 Lingulodinium Wall, p. 109.
Synopsis. A chorate gonyaulacacean (cribroperidinioid)
cyst with a subspherical central body. Acavate. The cen -
tral body is smooth or ornamented with features of low
relief and bears numerous simple, hollow, apparently
nontabular processes that are variable in length and
distal ending. Archaeopyle precingular, resulting from
loss of a variable number of precingular plates (one to
five), or epicystal. When the archaeopyle is precingular,
the opercular pieces are usually free and separate.
Remarks. We do not know if the variation in archaeopyle
development among specimens of Lingulodinium bears
any relationship to age or palaeoenvironment.
Lingulodinium funginum (Morgenroth 1966a)
Islam 1983a
(Plate 10, figs 1, 2)
1966a Baltisphaeridium funginum Morgenroth, p.
17, 18, plate 3, figs 7, 8.
1983a Lingulodinium funginum (Morgenroth) –
Islam, p. 341.
Age. Peak of Lingulodinium spp. inconsistent within
Eocene – it probably has greater palaeoenvironmental
than biostratigraphic significance.
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56
Remarks. According to Morgenroth (1966a), Lingulo -
dinium funginum is characterised by having a variable
number of processes that are distally bulbose, in
contrast to species such as Lingulodinium machaeropho-
rum that have acuminate process endings. Specimens of
Lingulodinium with bulbose process endings are com -
mon in some samples from Labrador Margin wells and
these are included here in Lingulodinium funginum.
However, there is a caveat. Kokinos & Anderson (1995),
in laboratory experiments, noted that the length and
nature of the processes of resting cysts of the extant
Lingulodinium polyedrum show considerable variation.
The processes of the cysts, more familiarly known to
palynologists as Lingulodinium machaerophorum, could
be distally acuminate, bulbose, smooth or with small
granules or spines; process length could vary up to a
maximum of 10–12 μm. Based on their findings, Ko -
kinos & Anderson (1995) considered Lingulodinium
funginum, Lingulodinium sadoense and Lingulodinium
brevi spinosum to be taxonomic junior synonyms of
Lingulodinium machaerophorum. Because morphological
types included here in Lingulodinium funginum occur
only in the Eocene in the study material, we propose to
retain the species as defined in Morgenroth (1966a) and
prefer to avoid the use of infraspecific ranks (e.g. variety).
Retention may also provide useful information on pa -
laeo environments.
Lingulodinium machaerophorum (Deflandre &
Cookson 1955) Wall 1967
(Plate 10, figs 3, 4)
1955 Hystrichosphaeridium machaerophorum De -
flandre & Cookson, p. 274, plate 9, figs 4, 8.
1961 Baltisphaeridium machaerophorum (Deflandre
& Cookson) – Gerlach, p. 191, 192.
1966 Cleistosphaeridium machaerophorum (De -
flandre & Cookson) – Davey et al.,
p. 170; combination not validly published.
1967 Lingulodinium machaerophorum (Deflandre &
Cookson) – Wall, p. 109.
Age. LO not confirmed in the study area, but globally
the species extends to the present day.
Genus Nyktericysta Bint 1986
Type. Bint 1986, plate 4, figs 1, 2, 5, 6, text-fig. 3A, B,
as Nyktericysta davisii.
1986 Nyktericysta Bint, p. 148, 149.
1986 Balmula Bint, p. 158.
1999 Quantouendinium Mao Shaozhi et al., p. 155,
156.
Remarks. Fensome et al. (2009, p. 46) emended the
generic diagnosis of Nyktericysta and considered the two
wall layers to be the endophragm and ectophragm, to
better facilitate comparison with closely similar genera.
The ectophragm is finely perforate. According to the
emendation, Nyktericysta always has one apical and two
antapical horns. In addition, one or two lateral equato-
rial horns commonly occur, and these may have pre-
and postcingular extensions. The archaeopyle is apical,
with the formula A(1–4´) and the operculum usually
remains attached. Fensome et al. (2009, p. 46) noted
that Vesperopsis differs from Nyktericysta in not having
an ectophragm, and that Endoceratium and Pseudo -
ceratium differ in having only an apical, postcingular
and antapical horn rather than two antapical horns.
Mao Shaozhi et al. (1999, p. 156) differentiated
Quantouendinium from Nyktericysta Bint 1986 and
Ves per opsis Bint 1986 on the number and nature of the
horns. Fensome et al. (2009, p. 46) in their emendation
of Nyktericysta stated that it always has two antapical
horns and commonly can have one or two lateral, equa-
torial horns. This indicates that the main distinction
between Quantouendinium and Nyktericysta is that the
former has one postcingular horn. However, from Mao
Shaozhi et al. (1999, fig. 4), the location of both poste-
rior horns appears to be antapical. We thus consider
Quantouendinium to be a taxonomic junior synonym of
Nyktericysta.
Nyktericysta davisii Bint 1986
(Plate 10, figs 5, 6)
1986 Nyktericysta davisii Bint, p. 149, 150, 152,
153, plate 4, figs 1–12; plate 8, figs 1–6; text-
figs 3A–C, 4A, B, 10A, B.
Age. LO: late Albian.
Remarks. Nyktericysta davisii is characterised by the pres-
ence of five horns, of which the two lateral equatorial
have pre- and postcingular extensions. The ectophragm
is perforate. Tabulation may be indicated by low sutural
ridges.
Nyktericysta dictyophora He Chengquan et al. 1992
(Plate 10, figs 7, 8)
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57
1992 Nyktericysta dictyophora He Chengquan et al.,
p. 184, 190, 191, plate 1, figs 1–9.
1992 Nyktericysta dictyophora subsp. circularis He
Chengquan et al., p. 185, 191, plate 1, figs 7–9.
1992 Nyktericysta fusiformis He Chengquan et al., p.
185, 191, 192, plate 1, fig. 16; plate 2, figs 1–3.
1999 Quantouendinium dictyophorum (He Cheng -
quan et al.) – Mao Shaozhi et al., p. 156.
Age. LO: Cenomanian.
Remarks. Mao Shaozhi et al. (1999, p. 157) considered
Nyktericysta dictyophora subsp. circularis to be a junior
synonym of Quantouendinium dictyophorum (that is,
with the autonym). They also considered Nyktericysta
fusiformis to be taxonomic junior synonyms of this
species.
Nyktericysta tripenta (Bint 1986) Fensome et al.
2009
(Plate 10, figs 11, 12)
1986 Balmula tripenta Bint, p. 158, 160, plate 6,
figs 9–17; plate 7, fig. 8; text-fig. 6A, B.
2009 Nyktericysta tripenta (Bint) – Fensome et al.,
p. 46.
Age. LO: Albian.
Remarks. Fensome et al. (2009, p. 46) noted that Nyk -
tericysta tripenta has a coarse autophragmal reticulum,
with muri being up to 10 μm wide.
Genus Odontochitina Deflandre 1937
Type. Deflandre 1937, plate 18 (also labelled plate 15),
fig. 8, as Odontochitina silicorum.
1937 Odontochitina Deflandre, p. 94.
Remarks. The synopsis provided for Odontochitina by
Fen some et al. (2009, p. 46) does not allow for the
inclusion of forms with abbreviated horns.
Odontochitina ancala Bint 1986
(Plate 10, fig. 14)
1986 Odontochitina ancala Bint, p. 139, 140, plate
1, figs 2–8; plate 7, figs 1, 2; text-fig. 2A.
Age. LO: Cenomanian.
Remarks. According to Bint (1986, p. 140), “Odonto -
chitina ancala differs from O. operculata by having an
elbow and cingular notch in the right lateral horn,
localised perforations about midway along the horns,
and an elongate ventral extension of the antapical peri-
coel.”
Odontochitina costata Alberti 1961
(Plate 10, fig. 15)
1961 Odontochitina costata Alberti, p. 31, plate 6,
figs 10–13.
1962 Odontochitina striatoperforata Cookson &
Eisenack, p. 490, plate 3, figs 14–19.
Age. LO: latest Campanian.
Remarks. Odontochitina costata was emended by Clarke
& Verdier (1967, p. 58, 59), who considered Odonto -
chitina striatoperforata to be intergradational with, and
a taxonomic junior synonym of, Odontochitina costata.
Odontochitina porifera Cookson 1956
(Plate 10, fig. 13)
1956 Odontochitina porifera Cookson, p. 188, plate
1, fig. 7.
Age. LO: Santonian.
Remarks. In the Scotian Margin wells, the LO of Odon -
tochitina porifera is also Santonian (Fensome et al.
2009, p. 47).
Genus Oligosphaeridium Davey & Williams 1966a
Type. White 1842, plate 4, fig. 11, as Xanthidium tubi -
ferum var. complex.
1966a Oligosphaeridium Davey & Williams, p. 70, 71.
Remarks. In their synopsis for Oligosphaeridium, Fen -
some et al. (2009, p. 47) stated: “processes more or less
equal in size and general shape.” There are two excep-
tions, the first and fourth apical (1´, 4´) are generally
slender compared to the other apicals, although all the
apicals tend to be slender.
Oligosphaeridium albertense (Pocock 1962) Davey
& Williams 1969
(Plate 10, fig. 17)
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58
1962 Hystrichosphaeridium albertense Pocock, p. 82,
plate 15, figs 226, 227.
1962 Hystrichosphaeridium irregulare Pocock, p. 82,
plate 15, figs 228, 229.
1964 Hystrichosphaeridium coelenteratum Tasch in
Tasch et al., p. 195, plate 2, fig. 11.
1964 Hystrichosphaeridium dispare Tasch in Tasch
et al., p. 195, plate 2, fig. 8
1964 Hystrichosphaeridium reniforme Tasch in
Tasch et al., p. 193, plate 2, fig. 6.
1969 Oligosphaeridium albertense (Pocock) – Davey
& Williams, p. 5.
Age. LO: early Cenomanian.
Remarks. In his emendation of Oligosphaeridium alber -
tense, Brideaux (1977, p. 27, 28) described the process-
es as hollow, flared to tubiform, with the open distal
margins being “variably secate, occasionally aculeate or
serrate; the distal third of some processes variably fene -
strate ….” Jansonius (1986, p. 213) described the holo-
type of Oligosphaeridium albertense, which he re- illustrated
(his plate 4, figs 4, 5), as having processes that were prox-
imally nearly cylindrical but widened, and were strongly
flared distally. The distal margins of the processes were
“occasionally scalloped, carrying numerous coarse to fine,
slender spinules.” Jansonius (1986) considered Hystricho -
s phaeridium (as Oligosphaeridium) irregulare to be a taxo-
nomic junior synonym of Oligosphaeridium albertense.
Stover & Evitt (1978, p. 68, 69) regarded Hystricho -
sphae ridium (as Oligosphaeridium) coelenteratum, Hy -
stri chos phaeridium (as Oligosphaeridium) dispare and
Hystrichosphaeridium (as Oligosphaeridium) reniforme as
taxonomic junior synonyms of Oligosphaeridium irre -
gulare. Thus, by implication, Oligosphaeridium coelen-
teratum, Oligosphaeridium dispare and Oligosphaeri-
dium reniforme are all taxonomic junior synonyms of
Oligo sphaeridium albertense.
Oligosphaeridium pulcherrimum (Deflandre &
Cookson 1955) Davey & Williams 1966a
(Plate 10, figs 18, 19)
1955 Hystrichosphaeridium pulcherrimum Deflandre
& Cookson, p. 270, 271, plate 1, fig. 8; text-
figs 21, 22.
1966a Oligosphaeridium pulcherrimum (Deflandre &
Cookson) – Davey & Williams, p. 75, 76.
Age. LO: regional – Santonian; LAD – Danian. Not
plotted.
Remarks. Included here in Oligosphaeridium pulcherri-
mum are those forms that have a mixture of perforated
and unperforated processes. Oligosphaeridium complex
has no perforated processes.
Oligosphaeridium totum Brideaux 1971
(Plate 10, fig. 20)
1971 Oligosphaeridium totum Brideaux, p. 88, 89,
plate 25, figs 53–55, 57.
1971 Oligosphaeridium diastema Singh, p. 337,
plate 55, figs 4, 5; plate 56, figs 1, 2.
Age. LO: early Cenomanian.
Remarks. Based on nannofossil control, Fensome et al.
(2009) considered the last occurrence for Oligo sphae r -
idium totum to be in the early Cenomanian.
Genus Operculodinium Wall 1967
Type. Deflandre & Cookson 1955, plate 8, figs 3, 4, as
Hystrichosphaeridium centrocarpum.
1967 Operculodinium Wall, p. 110, 111.
Remarks. Operculodinium is a chorate gonyaulacacean
(criboperidinioidean) cyst with a spheroidal to slightly
ovoidal central body with a reticulate wall. The central
body bears numerous nontabular to contabular process-
es. Processes are generally solid, of the same size in indi-
vidual species and distally bifid to aculeate. The
archae opyle is precingular, with the formula P3´´ ; the
operculum is free. Fensome et al. (2009, p. 48) dis -
cussed the differences between Operculodinium and the
closely similar genus Exochosphaeridium.
Operculodinium centrocarpum (Deflandre &
Cookson 1955) Wall 1967
(Plate 11, figs 5, 6)
1955 Hystrichosphaeridium centrocarpum Deflandre
& Cookson, p. 272, 273, plate 8, figs 3, 4.
1961 Baltisphaeridium centrocarpum (Deflandre &
Cookson) – Gerlach, p. 192,193.
1965 Cordosphaeridium centrocarpum (Deflandre &
Cookson) – de Coninck, p. 33.
1966a Cordosphaeridium tiara subsp. centrocarpum
(Deflandre & Cookson) – Morgenroth, p. 26.
1967 Operculodinium centrocarpum (Deflandre &
Cookson) – Wall, p. 111.
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59
1969 Cordosphaeridium? microtriainum subsp.
centrocarpum (Deflandre & Cookson) – de
Coninck, p. 32.
1978 Cleistosphaeridium centrocarpum (Deflandre &
Cookson) – Jiabo, p. 61.
1983 Operculodinium echigoense Matsuoka, p. 126,
plate 7, figs 1, 2a, b, 3–5, 8.
1987 Operculodinium? echigoense Matsuoka –
Mudie, p. 804.
Age. LO: Tortonian.
Genus Palaeocystodinium Alberti 1961
Type. Alberti 1961, plate 7, fig. 12, as Palaeo cysto di -
nium golzowense.
1961 Palaeocystodinium Alberti, p. 20.
1963 Cystodiniopsis Vozzhennikova, p. 185.
Remarks. In their emendation of Palaeocystodinium, Fen -
some et al. (2009, p. 48) stated that they are “Peri -
diniacean (deflandreoid) cysts that are fusiform in shape,
with single prominent pointed horns apically and antapi-
cally; the horns are generally long and there may be a short
accessory antapical horn.” Fensome et al. (2009) differen-
tiated Palaeo cystodinium from the genus Sval bardella
Manum 1960 on the nature of the horns distally; the
latter has bluntly rounded apical and antapical horns.
Palaeocystodinium bulliforme Ioannides 1986
(Plate 11, figs 1, 2)
1986 Palaeocystodinium bulliforme Ioannides, p. 31,
plate 17, figs 2–5.
Age. LO: Selandian.
Palaeocystodinium golzowense Alberti 1961
(Plate 10, fig. 10)
1961 Palaeocystodinium golzowense Alberti, p. 20,
plate 7, figs 10–12; plate 12, fig. 16.
Age. LO: late Tortonian (Late Miocene).
Palaeocystodinium teespinosum Fensome et al.
2009
(Plate 11, fig. 3)
2009 Palaeocystodinium teespinosum Fensome et al.,
p. 50, plate 8, figs m–p.
Age. LO: early Rupelian.
Remarks. Palaeocystodinium teespinosum differs from
Pa laeo cystodinium golzowense in having delicate T-
shaped spinelets that are about 2 μm long and are espe-
cially common on the apical and antapical horns.
Genus Palaeohystrichophora Deflandre 1935
Type. Deflandre 1935, plate 8, fig. 4, as Palaeo hy -
strichophora infusorioides.
1935 Palaeohystrichophora Deflandre, p. 230.
Remarks. In their remarks for this genus, Fensome et al.
(2009, p. 50) compared Palaeohystrichophora to Sub ti -
lisphaera Jain & Millepied 1973. Both genera lack an
obvious archaeopyle, but Harker (1979, p. 374, fig. 1)
observed a combination archaeopyle with the formula
3I3P(1–3a + 3–5´´) in Palaeohystrichophora infusorioides;
according to Harker, the operculum remains attached
along the posterior or cingular margin. Bujak & Davies
(1983, p. 62, text-fig. 4) observed an archaeopyle, which
they termed the transverse archaeopyle, in some species
assigned to Subtilisphaera. This archaeopyle resulted from
the development of sutures between the apicals and three
anterior intercalary plates and between the anterior lateral
margins of the intercalaries. The major observable differ-
ence between the two genera is the presence of processes
on the pericyst of Palaeohystrichophora. Forms with pro -
cesses that are presently included in Subtilisphaera should
probably be included in Palaeohystrichophora.
Palaeohystrichophora infusorioides Deflandre 1935
(Plate 11, fig. 4)
1935 Palaeohystrichophora infusorioides Deflandre,
p. 230, 231, plate 8, fig. 4.
1943 Palaeohystrichophora paucisetosa Deflandre, p.
507, 508, text-fig. 26.
Age. LO: Campanian.
Genus Palaeoperidinium Deflandre 1934 ex
Sarjeant 1967
Type. Ehrenberg 1838, plate 1, fig. 4, as Peridinium
pyrophorum.
1934 Palaeoperidinium Deflandre, p. 968; name
not validly published.
1963 Pentagonum Vozzhennikova, p. 183; name
not validly published.
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60
1967 Palaeoperidinium Deflandre ex Sarjeant, p.
246, 247.
1967 Pentagonum Vozzhennikova ex Vozzhen -
nikova, p. 106; name illegitimate.
1970 Astrocysta Davey, p. 359.
Remarks. The synopses of Palaeoperidinium provided
by Stover & Evitt (1978, p. 217) and Fensome et al.
(2009, p. 51) are very similar. In their remarks, Fen -
some et al. (2009) took into account the emendation of
Palaeoperidinium by Evitt et al. (1998, p. 46, 48), who
noted that the most commonly preserved wall of the
cyst is an exophragm, which is unusual in having been
formed outside but in contact with the exterior surface
of the theca.
Palaeoperidinium pyrophorum (Ehrenberg 1838 ex
Wetzel 1933b) Sarjeant 1967
(Plate 11, figs 7, 8)
1838 Peridinium pyrophorum Ehrenberg, plate 1,
figs 1, 4; name not validly published.
1933b Peridinium pyrophorum Ehrenberg ex Wetzel,
p. 164, 165.
1967 Palaeoperidinium pyrophorum (Ehrenberg ex
Wetzel) – Sarjeant, p. 246.
1967 Peridinium basilium Drugg, p. 13, plate 1,
figs 9–11; plate 9, fig. 1a, b.
1967 Pentagonum marginatum Vozzhennikova, p.
107, plate 46, figs 1, 3, 4, 6; generic name
illegitimate.
1967 Pentagonum sibiricum Vozzhennikova, p. 106,
107, plate 46, figs 2, 5.
1967 Peridinium conicum var. larjakiense Vozzhen -
nikova, p. 71, 72, plate 16, figs 1a, b, 2a, b.
1973 Palaeoperidinium deflandrei Lentin & Wil -
liams, p. 105.
1981 Palaeoperidinium larjakiense (Vozzhennikova)
– Lentin & Williams, p. 210.
Age. LO: Selandian; peak: early Danian.
Remarks. Gocht & Netzel (1976, p. 403–405) and Evitt
et al. (1998, p. 48, 49) provided comprehensive, concise
emendations of Palaeoperidinium pyrophorum, and re -
solved its relationship to the thecate equivalent.
Genus Palynodinium Gocht 1970
Type. Gocht 1970, fig. 4, nos 1a–c, as Palynodinium
grallator.
1970 Palynodinium Gocht, p. 135, 137, 138, 140.
Palynodinium grallator Gocht 1970
(Plate 11, figs 9, 10)
1970 Palynodinium grallator Gocht, p. 135, 137,
138, 140, fig. 2a–e; fig. 4, nos 1a–c, 2a, b, 3a,
b, 4a, b, 5a, b, 6a, b, 7, 8; fig. 5, nos 1, 2a, b.
Age. LO: latest Maastrichtian.
Genus Petalodinium Williams et al. 2015
Type. Williams & Downie 1966b, plate 20, figs 1, 2, as
Wetzeliella condylos.
2015 Petalodinium Williams et al., p. 307.
Remarks. Williams et al. (2015) erected the genus Peta -
lo dinium for wetzelielloidean dinocysts with a latiepeli-
form archaeopyle and a pericyst that is smooth or
or namented with features of low relief.
Petalodinium condylos (Williams & Downie
1966b) Williams et al. 2015
(Plate 11, figs 11, 12)
1966b Wetzeliella condylos Williams & Downie, p.
193, 194, plate 20, figs 1, 2.
1976 Rhombodinium condylos (Williams & Downie)
– Lentin & Williams, p. 128.
1979 Dracodinium condylos (Williams & Downie) –
Costa & Downie, p. 43.
2015 Petalodinium condylos (Williams & Downie) –
Williams et al., p. 308.
Age. LO: Ypresian.
Remarks. As Williams et al. (2015) have demonstrated,
the holotype of Petalodinium condylos (Williams &
Dow nie 1966b, plate 20, fig. 1, as Wetzeliella condylos),
which is the type of Petalodinium, has a latiepeliform
archaeopyle. Apparently related to the latiepeliform
archaeopyle is the nature of the apical horn, which in
Petalodinium condylos is reduced or absent, the apex
being rounded. The pericyst is verrucate to tuberculate.
Often, the intratabular ornamentation delineates the
tabulation: other verrucae or tubercles are penitabular.
Both peri- and endophragm are thick, about 3 μm.
Some specimens observed in the Labrador Margin
samples are devoid of ornamentation, but are otherwise
identical.
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61
Genus Phelodinium Stover & Evitt 1978
Type. Corradini 1973, plate 28, fig. 3, as Deflandrea
pentagonalis.
1978 Phelodinium Stover & Evitt, p. 117, 118.
Synopsis. A dorso-ventrally compressed, proximate
protoperi diniacean (protoperidinioidean) cyst with one
apical and two antapical horns. The cyst is pentagonal
with convex to concave lateral sides. It is produced into
one apical and two antapical horns that can be of vari-
able length, are usually acuminate distally, and may
have solid tips. Cornucavate where two wall layers can
be discerned. Cingulum sometimes present. Endocyst,
where observable, and pericyst smooth or ornamented
with features of low relief. Archaeopyle intercalary, with
formula I(2a), the second anterior intercalary plate is
deltaform; operculum usually free.
Remarks. The emendation of Phelodinium by Mao
Shaozhi & Norris (1988, p. 51, 52) related to the
archaeopyle index, given in the original diagnosis as 0.3
to 0.4 by Stover & Evitt (1978, p. 117). Mao Shaozhi
& Norris (1988) expanded the index to between 0.3
and 0.6; these authors also classified the archaeopyle as
standard hexa 2a. One difficulty with Phelodinium is
determining if it has one or two wall layers. According
to Stover & Evitt (1978, p. 118) “Phelodinium differs
from Lejeunia in being cavate and in having a peridinioid
outline with straight to concave sides. Lejeunia has an
autophragm only, and its lateral margins are normally
convex.” That there are two wall layers in Phelodi nium is
often extremely difficult to discern and the outline of the
cyst in the two genera appears to be variable.
Phelodinium kozlowskii (Górka 1963) Lindgren
1984
(Plate 12, figs 1–3)
1963 Lejeunia kozlowskii Górka, p. 41, plate 5, fig. 4.
1970 Astrocysta kozlowskii (Górka) – Davey, p. 369.
1977 Senegalinium kozlowskii (Górka) – Harland,
p. 189.
1984 Phelodinium kozlowskii (Górka) – Lindgren,
p. 181.
Age. LO: Danian.
Remarks. Harland (1973, p. 673) and Harker & Sarjeant
(1975, p. 223) considered Phelodinium kozlowskii to be a
taxonomic junior synonym of Phe lodinium tricuspe. In
their emendation of Phelodinium tricuspe, Lejeune-Car -
pentier & Sarjeant (1981, p. 20) stated that the “Phragma
[is] apparently composed of a single layer (autophragm).”
These authors also ob served that in the holotype of
Phelodinium (as Leje unecysta) tricuspe only one wall layer
could be discerned: this observation explains why these
authors accepted the transfer of the species to Lejeune -
cysta by Artzner & Dörhöfer (1978). Lindgren (1984, p.
181–183) recorded two wall layers in Phelo dinium
kozlowskii, with the endophragm being closely appressed
to the periphragm except at the tips of the horns. Some
of the specimens of Phelodinium kozlowskii encountered
in this study show a similar morphology, which is why
the species is retained in Phelodinium and separated from
Phelodinium tricuspe; assuming that the holotype of the
latter species has a single wall layer, it should be included
in Lejeunecysta.
Genus Phthanoperidinium Drugg & Loeblich Jr.
1967
Type. Drugg & Loeblich Jr. 1967, plate 1, fig. 4, as
Phtha noperidinium amoenum.
1967 Phthanoperidinium Drugg & Loeblich Jr.,
p. 182.
Remarks. Fensome et al. (2009, p. 54) provided a com -
prehensive synopsis of Phthanoperidinium, which in -
cluded the findings by Edwards & Bebout (1981, p.
36) and Islam (1982, p. 306) on the variability and
complexity of the archaeopyle. Although the variability
has not been noted rigorously, if the trend is similar to
that in other peridiniaceans, it is probable that the vari-
ous types have different stratigraphic ranges. This has
already been demonstrated in Phthanoperidinium gemi-
natum and Phthanoperidinium regale, whose only dif -
ference is in the nature of the archaeopyle.
Phthanoperidinium coreoides (Benedek 1972)
Lentin & Williams 1976
(Plate 11, figs 13, 14)
1972 Hystrichogonyaulax coreoides Benedek, p. 20,
plate 9, fig. 4a–c.
1976 Phthanoperidinium coreoides (Benedek) –
Lentin & Williams, p. 76.
1976 Phthanoperidinium tritonium Eaton, p. 299,
300, plate 17, figs 2, 3, 6, 7; text-figs 23C,
24A, B.
Age. LO: Rupelian.
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62
Remarks. Following Fensome et al. (2009, p. 55), Phtha -
no peridinium coreoides and Phthanoperidinium comatum
are separate species, distinguished on the relative length
of the processes. In Phthanoperidinium coreoides, the pro -
cesses are about one third the diameter of the central
body; in Phthanoperidinium comatum, the processes are
about one half the diameter of the central body.
Phthanoperidinium levimurum Bujak in Bujak et
al. 1980
(Plate 11, fig. 15)
1980 Phthanoperidinium levimurum Bujak in Bujak
et al., p. 74, plate 19, figs 13–16; text-figs
20E, 22B.
Age. LO: Priabonian.
Remarks. Phthanoperidinium levimurum is characterised
by having the tabulation delineated by membranes,
which are usually smooth distally but may sometimes be
denticulate.
Phthanoperidinium multispinum Bujak in Bujak et
al. 1980
(Plate 11, figs 16, 17)
1980 Phthanoperidinium multispinum Bujak in
Bujak et al., p. 74, plate 19, figs 17–19; text-
fig. 20F.
Age. LO: latest Priabonian.
Phthanoperidinium regale Bujak 1994
(Plate 11, figs 18, 19)
1994 Phthanoperidinium regale Bujak, p. 130, plate
4, figs 4–6.
Age. LO: Lutetian.
Remarks. Phthanoperidinium regale has ornamentation
similar to Phthanoperidinium geminatum Bujak in Bujak
et al. 1980, but differs in having a combination archae -
opyle, with the formula IP(2a+4´´). Phano per idinium gemi-
natum has the more usual archaeopyle type for the
genus, losing the 2a plate only. In Phthanoperidinium
regale, the operculum is free.
Phthanoperidinium stockmansii (de Coninck 1975)
Lentin & Williams 1977a
(Plate 11, fig. 20)
1975 Peridinium stockmansii de Coninck, p. 97, 98,
plate 17, figs 18–37.
1977a Phthanoperidinium stockmansii (de Coninck)
– Lentin & Williams, p. 131.
1976 Phthanoperidinium echinatum Eaton, p. 298,
299, plate 17, figs 8, 9, 12; text-fig. 23B.
1980 Phthanoperidinium? pseudoechinatum Bujak in
Bujak et al., p. 75, 76, plate 19, fig. 20; text-
fig. 20C.
Age. LO: Priabonian.
Remarks. Phthanoperidinium stockmansii has penitabular
rows of small processes that distally are clavate. Rows of
processes usually occur in parallel pairs, somewhat like
railway tracks. De Coninck (1977, p. 40) considered
Phtha noperidinium echinatum to be a junior taxonomic
synonym of Phthanoperidinium stockmansii. Islam (1982,
p. 315) agreed with this synonymy, but mistakenly
thought that Phthanoperidinium echinatum was the
senior name. Accepting Islam’s synonymy of Phthano -
peri dinium pseudoechinatum with Phthanoperi dinium
echinatum, the former thus becomes a junior taxonom-
ic synonym of Phtha no peridinium stockmansii.
Genus Piladinium Williams et al. 2015
Type. Michoux 1988, plate 1, figs 2, 3, as Kisselovia
columna.
2015 Piladinium Williams et al., p. 308, 309.
Remarks. Piladinium is a wetzelielloidean genus with a
latiepeliform archaeopyle and processes connected by an
ectophragmal membrane. For comparison with other
wetzelielloidean genera, see Williams et al. (2015).
Piladinium columna (Michoux 1988) Williams et
al. 2015
(Plate 12, figs 5, 6)
1988 Kisselevia columna Michoux, p. 28, 30, plate 1,
figs 2, 3, 5, 6; plate 2, figs 3–5; text-fig. 7A, B.
1989 Charlesdowniea columna (Michoux) – Lentin
& Vozzhennikova, p. 74.
2015 Piladinium columna (Michoux) – Williams et
al., p. 309.
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63
Age. LO: Ypresian.
Remarks. Several of the observed Labrador Margin
specimens of Piladinium columna have reduced num -
bers of processes on the mid-ventral and mid-dorsal
regions. The morphology of these specimens is thus
approaching that of Piladinium edwardsii.
Piladinium edwardsii (Wilson 1967b) Williams et
al. 2015
(Plate 12, figs 7, 8)
1967b Wetzeliella edwardsii Wilson, p. 477, figs 8, 9.
1978 Kisselevia edwardsii (Wilson) – Stover & Evitt,
p. 111.
1989 Charlesdowniea edwardsii (Wilson) – Lentin
& Vozzhennikovia, p. 227.
2015 Piladinium edwardsii (Wilson) – Williams et
al., p. 309.
Age. LO: early Ypresian. Not plotted.
Genus Pseudoceratium Gocht 1957
Type. Gocht 1957, plate 18, fig. 1, as Pseudoceratium
pelliferum.
1957 Pseudoceratium Gocht, p. 166.
1962 Eopseudoceratium Neale & Sarjeant, p. 446.
1966a Doidyx Sarjeant, p. 205.
Synopsis. A dorso-ventrally compressed, proximate cera-
tiacean cyst with single apical, postcingular and antapical
horns. Wall formed of one or two layers. If two-layered,
there can be endophragm and periphragm or endophragm
and ectophragm. Tabulation often indicated by ornamen-
tation if present. Ornamentation may be absent or it may
be granular or consist of short processes, which may be
trabeculate. Cingulum sometimes obvious. Archaeopyle
apical, with formula A(1–4´), operculum free; sulcal notch
offset to the left.
Pseudoceratium sp.
(Plate 12, fig. 4)
Remarks. Our specimens of Pseudoceratium differ from
Pseudoceratium pelliferum in having longer processes
and in generally lacking a lateral horn.
Genus Raphidodinium Deflandre 1936
Type. Deflandre 1936, plate 10, figs 1, 2, 7, Raphido -
dinium fucatum.
1936 Raphidodinium Deflandre, p. 184, 185.
Synopsis. Chorate cyst with ovoidal central body and a
clearly defined cingulum. The cingulum can subdivide
the central body into equal epi- and hypocysts or be ante-
riorly located, dividing the cyst into a short epicyst and
much longer hypocyst. Tabulation marked by mem -
branes or ridges and long, slender processes that are clus-
tered along the cingulum and to a lesser extent in the
antapical area. Processes are distally blunt to multifur-
cate. Archaeopyle indeterminate.
Remarks. Marheinecke (1992, p. 79) considered the
archaeopyle of Raphidodinium fucatum subsp. com pac -
tum Marheinecke 1992 to be possibly precingular.
Raphidodinium fucatum Deflandre 1936
(Plate 12, figs 9, 10)
1936 Raphidodinium fucatum Deflandre, p. 185,
186, plate 10, figs 1–7.
Age. LO: Campanian.
Remarks. The unequal sizes of the epi- and hypocysts
suggest that Raphidodinium fucatum could be a clado -
pyxiacean.
Genus Reticulosphaera Matsuoka 1983
Type. Matsuoka 1983, plate 4, fig. 8, as Reticulato -
sphaera stellata.
1983 Reticulatosphaera Matsuoka, p. 116.
Reticulatosphaera actinocoronata (Benedek 1972)
Bujak & Matsuoka 1986
(Plate 12, figs 11, 12)
1972 Cleistosphaeridium actinocoronatum Benedek,
p. 34, plate 12, fig. 13; text-fig. 12.
1978 Areosphaeridium? actinocoronatum (Benedek)
– Stover & Evitt, p. 20.
1983 Reticulatosphaera stellata Matsuoka, p. 116,
117, plate 4, figs 8–11; text-fig. 10.
1986 Reticulatosphaera actinocoronata (Benedek
1972) – emend. Bujak & Matsuoka 1986,
p. 238.
Age. LO: earliest Zanclian. Not plotted.
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64
Genus Rhombodinium Gocht 1955 emend.
Williams, Damassa, Fensome & Guerstein in
Fensome et al. 2009
Type. Gocht 1955, text-fig. 1c, as Rhombodinium draco.
1955 Rhombodinium Gocht, p. 85.
1961 Wetzeliella subgenus Rhombodinium (Gocht)
– Alberti, p. 9.
2009 Rhombodinium Gocht 1955 emend. Wil-
liams, Damassa, Fensome & Guerstein in
Fensome et al., p. 57.
Remarks. We fully concur with the emendation in
Fensome et al. (2009), which emphasises the generic
significance of the archaeopyle. According to the emend-
ed diagnosis, species of Rhombodinium must have a
solei form archaeopyle, which always has an operculum
that is attached apically. Rhombodinium was retained at
generic rank by Lentin & Williams (1977a, p. 139).
Rhombodinium draco Gocht 1955
(Plate 12, figs 13, 14)
1955 Rhombodinium draco Gocht, p. 86, text-fig.
1a–c.
1961 Wetzeliella subgenus Rhombodinium draco
(Gocht) – Alberti, p. 8.
Age. LO: Priabonian.
Remarks. Vozzhennikova (1967, p. 168) retained this
taxon in Rhombodinium.
Rhombodinium porosum Bujak 1979
1979 Rhombodinium porosum Bujak, p. 314, 315,
plate 1, figs 3, 5–8; plate 2, fig. 11; text-fig. 8C.
Age. LO: Bartonian.
Genus Rottnestia Cookson & Eisenack 1961a
Type. Eisenack 1954, plate 9, fig. 5, as Hystrichosphaera
borussica.
1961a Rottnestia Cookson & Eisenack, p. 40, 42.
Rottnestia borussica (Eisenack 1954) Cookson &
Eisenack 1961a
(Plate 13, fig. 1)
1954 Hystrichosphaera borussica Eisenack, p. 62,
plate 9, figs 5a, b, 6, 7.
1961a Rottnestia borussica (Eisenack) – Cookson &
Eisenack, p. 42.
1966a Triblastula borussica (Eisenack) – Morgenroth,
p. 15, 16.
1966b Hystrichosphaeropsis borussica (Eisenack) –
Sarjeant, p. 139; combination not validly
published.
Age. LO: Lutetian. Not plotted.
Genus Scalenodinium gen. nov.
Type. Plate 12, fig. 15, as Scalenodinium scalatum.
Etymology. The name is from the Latin scalenus, mean-
ing unequal, uneven, odd, in reference to the unequal
variation in apical and antapical development.
Description. Pericoel elongate, with an apical horn that
is acuminate to rounded distally and generally a round-
ed antapex, although the latter may be extended into a
short protuberance. When the endocyst is present, the
cyst is bicavate. Cingulum present or faintly expressed.
Periarchaeopyle intercalary, generally with formula I2a,
and with a free operculum. The hexa 2a plate is steno -
deltaform. In some specimens, the apparent loss of the
apical area (Plate 12, figs 16–19) or of other intercalary
plates besides the 2a (Plate 12, fig. 15) indicates a com -
pound and/or combination archaeopyle.
Remarks. Scalenodinium differs from Isabelidinium in
having a stenodeltaform, rather than a lati- to iso-ome ga -
form 2a plate. Palaeocystodinium, which like Scaleno -
dinium has a stenodeltaform 2a plate, differs from the
latter genus in always having more or less apical and
antapical horns and a periarchaeopyle that is always
formed from the loss of a single plate, the 2a plate,
rather than several plates as in Scalenodinium.
Scalenodinium scalenum sp. nov.
(Plate 12, figs 15–20)
Holotype. Plate 12, fig. 15, from a cuttings sample at
2135–2145 m in Gilbert F-53, GSC type collection no.
137916, sample P39484, slide 01, co-ordinates 19.3 ×
106.5, England Finder T37/4. Pericyst length 81 μm,
width 31 μm. The age determined for the sample from
which the holotype was recovered is Ypresian.
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Etymology. The name is from the Latin scalenus, mean-
ing unequal, uneven, odd, in reference to the unequal
variation in apical and antapical development.
Description. A species of Scalenodinium with an elon-
gate pericoel that has an apex bearing a well-developed,
distally acuminate to rounded apical horn and an
antapex that is generally rounded, though an antapical
protuberance may occasionally be developed. When an
endocyst is present, the cyst is bicavate. The pericyst is
verrucate to granulate in the mid-dorsal and mid-
ventral regions. A cingulum is rarely developed. Both
pericyst and endocyst have walls that are, at the most, 1
μm thick. Periarchaeopyle intercalary, generally with
formula I2a, operculum free. The hexa 2a plate is sten-
odeltaform. In some specimens, the apparent loss of the
apical area (Plate 12, figs 16–19) or of other intercalary
plates besides the 2a (Plate 12, fig. 15) indicates a
compound and/or combination archaeopyle.
Size. Pericyst length 77–86 μm, pericyst width 38–56
μm, apical horn length 18–27 μm, apical horn maxi-
mum breadth 11–14 μm.
Age. LO: Ypresian.
Remarks. Scalenodinium scalenum shows some variation
in the antapical region: although generally rounded ant -
apically, some specimens have a prominent antapical
horn that is more or less centrally located. The pericyst
invariably has folds running across its surface. It is diffi-
cult to be specific about the nature of the archaeopyle
since there is a common tendency for the apical polar
area to break up. Thus this breakup could be interpreted
to denote a compound or combination archaeopyle.
Width and shape of the distal terminations of the horn
or horns vary considerably.
Genus Schematophora Deflandre & Cookson 1955
Type. Deflandre & Cookson 1955, plate 6, figs 11, 12,
as Schematophora speciosa.
1954 Schematophora Deflandre & Cookson, p.
1237 (name not validly published).
1955 Schematophora Deflandre & Cookson, p. 262.
Remarks. The synopsis provided by Fensome et al.
(2009) covers all the salient points regarding the mor -
phology of Schematophora.
Schematophora speciosa Deflandre & Cookson
1955
(Plate 13, fig. 2)
1955 Schematophora speciosa Deflandre & Cookson,
p. 262, 263, plate 6, figs 11–13; plate 7, fig. 11.
Age. LO: Priabonian.
Genus Senegalinium Jain & Millepied 1973
Type. Jain & Millepied 1973, plate 1, figs 1–3, as
Senegalinium bicavatum.
1973 Senegalinium Jain & Millepied, p. 22, 23.
Synopsis. Peridiniacean (deflandreoid) cysts with a peri-
dinioid, usually elongate pericyst and two antapical
horns that are more or less of equal length. Bicavate
with a circular to pentagonal endocyst. Pericyst surface
smooth or with low ornament, sometimes with cingu-
lum indicated. Periarchaeopyle intercalary, resulting
from the loss of the iso- to stenodeltaform deltaform 2a
plate; the operculum commonly remains attached
along the posterior margin.
Remarks. Alterbidinium differs from Senegalinium in
having two antapical horns that are of unequal length.
Since Alterbidinium contains species that may have a
combination intercalary–precingular archaeopyle result-
ing from the loss of the 2a and 4´´ plate, it would not be
surprising to find the same variability in Senegalinium.
Senegalinium iterlaaense Nøhr-Hansen &
Heilmann-Clausen 2001
(Plate 13, figs 3, 4)
2001 Senegalinium iterlaaense Nøhr-Hansen &
Heilmann-Clausen, p. 164, 166–168, fig. 6,
nos 1–6.
Age. LO: Selandian.
Remarks. Senegalinium iterlaaense differs from Isabelidi -
nium viborgense in having a striate periphragm.
Genus Senoniasphaera Clarke & Verdier 1967
Type. Clarke & Verdier 1967, plate 14, fig. 8, as Seno -
niasphaera protrusa.
1967 Senoniasphaera Clarke & Verdier, p. 61.
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66
Remarks. The cavate cysts included in Senoniasphaera
have the typical offset sulcal notch of areoligeraceans.
The operculum of the apical archaeopyle is always free.
Senoniasphaera inornata (Drugg 1970) Stover &
Evitt 1978
(Plate 13, fig. 5)
1970 Chiropteridium inornatum Drugg, p. 811,
812, fig. 3C–F.
1978 Senoniasphaera inornata (Drugg) – Stover &
Evitt, p. 80.
Age. LO: Danian.
Remarks. Brinkhuis & Schiøler (1996) recorded Seno -
nias phaera inornata from the early Danian of the Geul -
hemmerberg Cretaceous–Palaeogene boundary section
in Limburg, south-eastern Netherlands. This observa-
tion accords with the age of this species in sections at
Stevns Klint (Denmark) and Alabama as recorded by
Hansen et al. (1986) and Habib (1994) respectively.
Senoniasphaera inornata therefore appears to be a good
Danian index species. Williams et al. (2004) placed the
LO of the species close to the top of the Danian.
Senoniasphaera microreticulata Brideaux &
McIntyre 1975
(Plate 13, fig. 6)
1975 Senoniasphaera microreticulata Brideaux &
McIntyre, p. 35, plate 11, figs 7–12; plate 12,
figs 1–8.
1981 Canningia microreticulata (Brideaux &
McIntyre) – Below, p. 31.
Age. LO: Cenomanian.
Remarks. Lentin & Williams (1981, p. 33) retained this
species in Senoniasphaera.
Senoniasphaera rotundata Clarke & Verdier 1967
(Plate 13, fig. 7)
1967 Senoniasphaera rotundata Clarke & Verdier,
p. 62, 63, plate 14, figs 1–3; text-fig. 25.
Age. LO: latest Campanian.
Remarks. In Senoniasphaera rotundata, the endocyst does
not protrude into the antapical horns.
Genus Simplicidinium gen. nov.
Type. Eaton 1976, plate 21, fig. 5, as Impletosphae ri -
dium insolitum.
Etymology. The name derives from the Latin simplicis,
meaning simplicity, in reference to the relatively simple
morphology of the cyst, comprising a spiny ball with an
apical archaeopyle.
Diagnosis. Proximochorate to chorate dinoflagellate cysts
with a more or less symmetrical spheroidal to ovoidal
central body. Spines or processes numerous, isolated and
non-tabulate, distally closed, with symmetrical distal
terminations. Archaeopyle apical, type 4A1´– 4´, opercu-
lum attached or detached.
Remarks. There seems to be considerable confusion
currently surrounding the generic assignment of chorate
species with non-tabulate spines/processes and a cryptic,
or not clearly discernible or consistent archaeopyle.
Forms with a consistent apical archaeopyle and at least
some asymmetrical processes are assignable to Cleisto -
sphaeridium (Eaton et al. 2001). Islam (1993) proposed
Downiesphaeridium, purportedly for forms with an apical
archaeopyle and simple non-tabulate processes. The
processes of the type of Downies phae ridium are typical of
Lin gulodinium, and the archaeopyle in the type of Dow -
nies phae ridium looks precingular; it is thus suggested here
that Downies phae ridium may be a taxonomic junior
synonym of Lingu lo dinium, but at least should not be
used beyond the type. Many authors have assigned
species that they consider to be chorate dinoflagellates
but show no evidence of an archaeopyle to Impleto -
sphaeridium, the type of that genus being suitably cryptic
in its archaeopyle type. Similar species deemed not to be
dinoflagellates are as sign able to the acritarch genus Balti -
sphaeridium.
Simplicidinium insolitum (Eaton 1976) comb. nov.
(Plate 13, figs 9, 10)
1976 Impletosphaeridium insolitum Eaton, p. 308,
plate 21, figs 5, 8; text-fig. 25B.
1978 Cleistosphaeridium? insolitum (Eaton) – Stover
& Evitt, p. 31.
Age. Local peak: earliest Bartonian.
Remarks. Eaton (1976, p. 308) commented on the pres-
ence of a polygonal opening in some of the specimens of
Simplicidinium (as Impletosphaeridium) insolitum; such
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67
openings are readily apparent in his ac companying illu -
strations (Eaton 1976, plate 21, figs 5, 8). We thus assign
this species, as type, to Simplicidinium on the basis of the
morphology of the holotype and because several of the
specimens in this study also have an apical archaeopyle.
Genus Sophismatia Williams et al. 2015
Type. Williams & Downie 1966a, plate 20, figs 2, 4,
text-fig. 49, as Wetzeliella tenuivirgula.
2015 Sophismatia Williams et al., p. 312, 313.
Remarks. Williams et al. (2015) are followed here in re -
stricting Sophismatia to wetzelielloideans with an equie-
peliform 2a archaeopyle and trabeculate processes.
Sophismatia tenuivirgula (Williams & Downie
1966b) Williams et al. 2015
(Plate 13, fig. 8)
1966b Wetzeliella tenuivirgula Williams & Downie,
p. 188, 189, plate 19, figs 1, 2, 4, 5, 7; text-
figs 49, 50.
1976 Kisselevia tenuivirgula (Williams & Downie) –
Lentin & Williams, p. 136.
1989 Charlesdowniea tenuivirgula (Williams &
Downie) – Lentin & Vozzhennikova, p. 227.
2015 Sophismatia tenuivirgula (Williams &
Downie) – Williams et al., p. 313.
Age. LO: Lutetian.
Remarks. The holotype of Sophismatia tenuivirgula (Wil -
liams & Downie 1966b, plate 19, figs 2–4; text-fig. 49)
clearly shows an equiepeliform archaeopyle, with the
endo- and periarchaeopyle being of almost identical size.
The only difference is anteriorly, where the periar-
chaeopyle extends slightly beyond the endoarchaeopyle.
Genus Spinidinium Cookson & Eisenack 1962
Type. Cookson & Eisenack 1962, plate 1, figs 1, 2, as
Spinidinium styloniferum.
1962 Spinidinium Cookson & Eisenack, p. 489.
2003 Magallanesium Quattrocchio & Sarjeant, p.
138, 140.
2003 Volkheimeridium Quattrocchio & Sarjeant,
p. 136, 138.
Remarks. In their extensive review of Spinidinium, Voz z -
hennikovia and related genera, Sluijs et al. (2009) emended
the diagnosis of Spinidinium. They restricted the genus to
peridiniacean taxa with proximosutural spines, a steno- to
isodeltaform 2a plate and an I2a archaeopyle, with the
operculum typically attached posteriorly.
Spinidinium echinoideum (Cookson & Eisenack
1960a) Lentin & Williams 1976
(Plate 13, figs 11, 12)
1960a Deflandrea echinoidea Cookson & Eisenack,
p. 2, plate 1, figs 5, 6.
1976 Spinidinium echinoideum (Cookson &
Eisenack) – Lentin & Williams, p. 64.
1978 Vozzhennikovia echinoideum (Cookson &
Eisenack) – Stover & Evitt, p. 130.
2003 Spinidinium? echinoideum (Cookson &
Eisenack) – Quattrocchio & Sarjeant, p. 136.
Age. LO: Selandian.
Remarks. Sluijs et al. (2009, p. 47) noted: “Both the
holo type (Cookson & Eisenack 1960a, plate 1, Fig. 5)
and specimens illustrated in Sverdlove & Habib (1974,
plate 1, figs 3, 5, 6; text–fig. 1) show the 2a plate is iso -
deltaform to isothetaform. The 2a also remains at -
tached to the 4´´ plate. Further the ornamentation is
predominantly proximosutural. For the above reasons we
include this species in Spinidinium without question.”
Genus Spiniferites Mantell 1850
Type. Ehrenberg 1838, plate 1, fig. 5, as Xanthidium
ramosum, designated by Davey & Williams (1966b, p.
32), as lectotype of Hystrichosphaera ramosa.
1850 Spiniferites Mantell, p. 191.
1933b Hystrichosphaera Wetzel, p. 33; name not
validly published.
1937 Hystrichosphaera Wetzel ex Deflandre, p. 61.
1953 Hystrichokibotium Klumpp, p. 387.
Remarks. Spiniferites is characterised by having its gony -
aula coidean tabulation expressed by sutural ridges or
membranes as well as gonal and sometimes sutural
processes. Distally, the gonal processes are always trifur-
cate, and the sutural processes are always bifurcate. The
archaeopyle is precingular, with formula P3´´, with a
free operculum.
Spiniferites ovatus Matsuoka 1983
(Plate 13, fig. 13)
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1983 Spiniferites ovatus Matsuoka, p. 134,
135, plate 3, figs 1a–c, 2, 3a, b, 4a, b;
text-fig. 19A, B.
non 1984 Spiniferites ovatus Bujak, p. 192, plate 3,
figs 15–18; illegitimate junior homonym.
Age. LO: Messinian.
Remarks. Spiniferites ovatus differs from Spiniferites pseu-
dofurcatus in having much shorter processes with shorter
terminations.
Spiniferites pseudofurcatus (Klumpp 1953)
Sarjeant 1970
(Plate 13, fig. 14)
1953 Hystrichokibotium pseudofurcatum Klumpp, p.
388, plate 16, figs 12, 14.
1960 Hystrichosphaera tertiaria Eisenack & Gocht,
p. 515; text-fig. 4.
1966b Hystrichosphaera buccina Davey & Williams,
p. 42, 43, plate 4, fig. 1; text-figs 10, 11.
1969 Hystrichosphaera pseudofurcata (Klumpp) –
Gocht, p. 32.
1970 Spiniferites pseudofurcatus (Klumpp) –
Sarjeant, p. 76.
Age. LO in present study: late Serravalian. LO accord-
ing to Piasecki (2003): Tortonian.
Remarks. Spiniferites pseudofurcatus differs from Acho -
mo s phaera alcicornu only in having the tabulation clear-
ly ex pressed by ridges. Although so similar, the
stratigraphic ranges of the two species differ consider-
ably, with Spini ferites pseudofurcatus having a much
younger LO.
Spiniferites scabrosus (Clarke & Verdier 1967)
Lentin & Williams 1975
(Plate 13, fig. 15)
1967 Hystrichosphaera scabrosus Clarke & Verdier,
p. 49, 50, plate 9, figs 7–10; text-fig. 21.
1975 Spiniferites scabrosus (Clarke & Verdier) –
Lentin & Williams, p. 2155.
Age. LO: Maastrichtian.
Remarks. The central body of Spiniferites scabrosus has a
distinctive scabrate to granulate wall and bears long, slen-
der processes. In some specimens, the sutural ridges or
membranes are not obvious, but such forms are retained
here in Spiniferites scabrosus.
Genus Spongodinium Deflandre 1936
Type. Ehrenberg 1838, plate 1, figs 1, 6, as Peridinium
delitiense.
1936 Spongodinium Deflandre, p. 169, 170.
Synopsis. Gonyaulacacean (cribroperidinioid) cysts that are
proximate, spheroidal to subpolyhedral, with a round ed
antapex and usually an apical protrusion or horn. Acavate.
The wall is complexly reticulate to vesi culate, and its total
thickness is sometimes greater along the cingulum and at
the poles. Wall atabulate or with structure/ornament ar -
ranged to reflect or suggest tabulation. Archaeopyle precin-
gular, with formula P3´´, oper culum free.
Remarks. The synopsis is similar to that provided by
Fensome et al. (2009, p. 60). An ectophragm is not
developed but may be partially simulated on the cyst
outline by the outer margins of membranes that consti-
tute the wall. The genus Samlandia is very similar in
appearance to Spongodinium, but the precise morphol-
ogy of the type is not clear from available illustrations
(Eisenack 1954, plate 11, figs 12‒15). However, an
SEM micrograph of a specimen identified as Samlandia
chlamydophora by one of the authors shows that an
occasionally perforate ectophragm connects processes
in all but the sutural areas. Such a morphology supports
our interpretation that Samlandia can thus be differen-
tiated from Spongodinium.
Spongodinium delitiense (Ehrenberg 1838)
Deflandre 1936
(Plate 13, figs 16, 20)
1838 Peridinium delitiense Ehrenberg, p. 110, plate
1, figs 1, 6.
1936 Spongodinium delitiense (Ehrenberg) –
Deflandre, p. 170, 171.
Age. LO: early Danian.
Remarks. The Labrador Margin specimens of Spongo di -
nium delitiense show considerable variation in size. The
larger specimens have an LO in the Maastrichtian and
the smaller ones range up into the Danian.
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69
Spongodinium grossum (Manum & Cookson 1964)
comb. nov.
(Plate 14, figs 1–4)
1964 Chlamydophorella grossa Manum & Cookson,
p. 17, 18, plate 5, figs 1, 2.
1986 Chlamydophorella? grossa Ioannides, p. 16.
Age. LO: Campanian.
Remarks. In a description of Spongodinium (as Chlamy -
do pho rella) grossum, Ioannides (1986, p. 16) stated:
“Archae opyle of dubious position, possibly precingular,
type P (or 2P). Paracingulum often indicated by aligned
processes, which may be joined proximally….” It is
confirmed here that the archaeopyle is precingular,
resulting from the loss of the 3´´ plate. The operculum
is free. Chlamydophorella has an apical archaeopyle, so
this species is transferred to Spongodinium.
M. Pearce (personal communication 2015) has drawn
our attention to the similarity between Spongo dinium
grossum and Isabelidinium? extremum. The latter species
has a precingular archaeopyle and an outer wall or
periphragm that forms an apical horn and a discontin-
uous pericyst. Thus, this species should be retained as
Spongodinium? extremum, a combination originally pro -
posed by Lentin & Williams (1976). Spongodinium
grossum and Spongodinium? extremum may be synony-
mous; the only obvious difference seems to be in the
nature of the pericyst, which is more continuous and
less perforate in Spongodinium grossum.
Spongodinium obscurum (Manum & Cookson
1964) comb. nov.
(Plate 13, fig. 19)
1964 Scriniodinium obscurum Manum & Cookson,
p. 21, 22, plate 4, figs 5, 6.
1978 Scriniodinium? obscurum (Manum &
Cookson) – Stover & Evitt, p. 188.
2003 Endoscrinium obscurum (Manum & Cookson)
– Riding & Fensome, p. 23.
Age. LO: Santonian.
Remarks. As demonstrated by Ioannides (1986, plates
5–8), intergradation exists between Spongodinium (as
Endoscrinium) obscurum, Spongodinium grossum (as Chla -
my dophorella grossa) and a taxon that he called Spon go d -
inium sp. In their description of Endoscrinium
obscurum, Manum & Cookson (1964, p. 21) stated:
“There are no distinct connections running between
the capsule and the theca, but fine, irregular lines, the
true nature of which is obscured by the many folds, may
possibly represent supporting fibrils. The wall of the
capsule is up to 1 μm thick and sometimes indistinctly
dotted.” The comments of Manum & Cookson (1964)
and Ioannides (1986), together with observations in
this study, indicate that the species should be assigned
to Spongodinium, and that the wall is best considered a
complex autophragm, rather than being holocavate.
Genus Stichodinium Williams et al. 2015
Type. He Chengquan & Wang Kede 1990, plate 2, fig.
3, as Wilsonidium subtile.
2015 Stichodinium Williams et al., p. 314.
Remarks. Williams et al. (2015) erected the genus Sticho -
dinium for wetzelielloidean cysts with a latiepeliform
archaeopyle and sutural or penitabular ornamentation
that may be features of low relief or processes that are
distally free.
Stichodinium lineidentatum (Deflandre &
Cookson 1955) Williams et al. 2015
(Plate 13, figs 17, 18)
1955 Wetzeliella lineidentata Deflandre &
Cookson, p. 253, 254, plate 5, fig. 5; text-figs
17, 18.
1976 Wilsonidium lineidentatum (Deflandre &
Cookson) – Lentin & Williams, p. 139.
2015 Stichodinium lineidentatum (Deflandre &
Cookson) – Williams et al., p. 314.
Age. LO: Lutetian.
Remarks. The holotype of Stichodinium lineidentatum
(Deflandre & Cookson 1955, plate 5, fig. 5) clearly
shows a latiepeliform archaeopyle.
Genus Subtilisphaera Jain & Millepied 1973
Type. Jain & Millepied 1973, plate 3, fig. 31, as Sub -
tilisphaera senegalensis.
1973 Subtilisphaera Jain & Millepied, p. 26, 27.
Remarks. Our concept of the genus Subtilisphaera adheres
to the synopsis presented in Fensome et al. (2009), who
acknowledged uncertainty in interpreting the archaeopyle.
Lentin & Williams (1976, p. 118) considered that, where
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observable, Subtilisphaera has a combination archaeopyle,
with the formula AIP(3´+1–3a+3–5´´), the operculum remain-
ing attached along the posterior margin. Stover & Evitt
(1978, p. 238) considered the archaeopyle of Subtili -
sphaera to be “presumably intercalary”. Bujak & Davies
(1983, p. 62) observed specimens from the Early Cre -
taceous of offshore eastern Canada, which they attrib-
uted to Subtilisphaera, with complete or incomplete
archaeopyle sutures between the following plates:
2´/1a, 3´/1a, 3´/2a, 3´/3a. 4´/3a and with complete or
incomplete archaeopyle sutures between individual
intercalary plates. These authors termed this a trans-
verse archaeopyle.
Subtilisphaera perlucida (Alberti 1959) Jain &
Millepied 1973
(Plate 14, fig. 5)
1959 Deflandrea perlucida Alberti, p. 102, plate 9,
figs 16, 17.
1959 Deflandrea pirnaensis Alberti, p. 100, plate 8,
figs 1, 5.
1960 Scriniodinium cooksoniae Anderson, p. 30,
plate 9, figs 1–3.
1973 Subtilisphaera perlucida (Alberti) – Jain &
Millepied, p. 27.
1973 Subtilisphaera pirnaensis (Alberti) – Jain &
Millepied, p. 27.
1990 Subtilisphaera? pirnaensis (Alberti) – Harker &
Sarjeant in Harker et al., p. 133.
Age. LO: Albian.
Remarks. Fensome et al. (2009, p. 61, 62) considered
Subtilisphaera pirnaensis to be a taxonomic junior syno -
nym of Subtilisphaera perlucida.
Genus Surculosphaeridium Davey et al. 1966
emend. nov.
Type. Sarjeant 1960, plate 6, fig. 2, as Hystricho -
sphaeridium cribrotubiferum.
1966 Surculosphaeridium Davey et al., p. 160, 161.
Emended diagnosis. Chorate gonyaulacacean cysts with
spheroidal central bodies. Acavate. Processes are solid
and branched and/or distally furcate. There may be one
to four processes per plate, sometimes forming com -
plexes in cases where there is more than one process per
plate. Archaeopyle apical, with the formula A(1–4´);
operculum free.
Remarks. Surculosphaeridium is emended to allow
inclusion of forms with more than one process per
plate. However, there must be processes or process
complexes on all the plates, including the precin -
gulars.
Surculosphaeridium convocatum sp. nov.
(Plate 14, figs 6–8)
Holotype. Plate 14, fig. 7, from a cuttings sample at
3510–3520 m in South Labrador N-19, GSC type
collection no. 138128, sample P39834, slide 01, co-
ordinates 7.8 × 99.1, England Finder G29/3‒H28/2.
Central body length (without apical operculum) 30
μm, central body width 38 μm, processes up to about
12 μm. The sample from which the holotype derives is
dated as Barremian‒Aptian, indicating that the speci-
men represents caving.
Etymology. The epithet is from the Latin convocatum, to
call together or assemble, in reference to the grouping
of several processes per plate common in this species.
Diagnosis. A species of Surculosphaeridium in which at
least the larger plates have more than one and up to four
processes per plate. Processes may branch along their
length and are always furcate distally.
Size. Central body length (without apical operculum)
29‒34 μm, central body width 30‒38 μm, processes up
to about 15 μm; three specimens measured.
Age. LO: early Campanian. Not plotted.
Remarks. Some plates, especially smaller ones such as
cingulars, sulcals and apicals, may have only one process
per plate, but others have two to four. Processes are
characteristically furcate distally. Surculosphaeri dium
lon gi furcatum has one process per plate.
Genus Talladinium Williams, Damassa, Fensome
& Guerstein in Fensome et al. 2009
Type. Mao Shaozi & Norris 1988, plate 13, fig. 6, as
Charlesdowniea wulagense.
2009 Talladinium Williams, Damassa, Fensome &
Guerstein in Fensome et al., p. 61, 62.
Remarks. Williams, Damassa, Fensome & Guerstein in
Fensome et al. (2009) erected the genus Talladinium
for wetzelielloidean cysts with a soleiform archaeopyle
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71
and processes that are distally united by ectophragmal
membranes that delineate the tabulation.
Talladinium? clathratum (Eisenack 1938)
Williams, Damassa, Fensome & Guerstein in
Fensome et al. 2009
(Plate 14, figs 9, 10)
1938 Wetzeliella clathrata Eisenack, p. 187; text-fig. 5.
1976 Kisselevia? clathrata (Eisenack) – Lentin &
Williams, p. 136.
1989 Charlesdowniea clathrata (Eisenack) – Lentin
& Vozzhennikova, p. 227.
2009 Talladinium? clathratum (Eisenack) –
Williams, Damassa, Fensome & Guerstein in
Fensome et al., p. 62.
Age. LO: latest Bartonian. Not plotted.
Remarks. As noted in Fensome et al. (2009, p. 62), the
nature of the peri- and endoarchaeopyles in Talladi -
nium? clathratum can not be ascertained from the orig-
inal description of Eisenack (1938) or the expanded
description of Eisenack (1954). From the late Eocene to
early Oligocene age of the type material, however, it
seems reasonable to deduce that this species has a
soleiform archaeopyle. This explains why the species is
questionably included in Talladinium.
Talladinium pellis sp. nov.
(Plate 14, figs 11, 12, 16)
Holotype. Plate 14, figs 11, 12, from a cuttings sample
at 1840 m in Gjoa O-37, GSC type collection no.
138082, sample YD16082, slide 03, coordinates 050 ×
0907, England Finder E20/3. Size: pericyst length 94
μm, width 76 μm; endocyst length 61 μm, width 57
μm. The age determined for the sample from which the
holotype was recovered is late Ypresian.
Etymology. The epithet is from the Latin pellis, meaning
skin, in reference to the ectophragm, which surrounds
most of the pericyst. It is a noun in apposition.
Diagnosis. A species of Talladinium in which processes
of individual plates are distally united by a membrane
that mimics the outline of the underlying plate. The
ectophragm may be irregularly or regularly perforate.
Size. Pericyst length 89‒94 μm, width 76‒81 μm; endo-
cyst length 61 μm (both specimens), width 57‒64 μm;
two specimens measured.
Age. LO: Priabonian.
Remarks. The ectophragmal membranes in Talladi nium
pellis form linear complexes that mirror the outline of
the reflected plates, rather than forming a shield-like
covering as in Talladinium? clathratum. This difference
is not always easy to determine. Charlesdowniea coleothryp-
ta has identical ornamentation but differs in having an
equiepeliform archaeopyle.
Genus Tanyosphaeridium Davey & Williams 1966a
Type. Davey & Williams 1966a, plate 6, fig. 7, text-fig.
20, as Tanyosphaeridium variecalamum.
1966a Tanyosphaeridium Davey & Williams, p. 98.
Remarks. Tanyosphaeridium is characterised by an elon-
gate ellipsoidal central body, slender open processes and
an apical archaeopyle. The processes number about 30
or more and cannot be readily related to tabulation.
Tanyosphaeridium xanthiopyxides (Wetzel 1933a
ex Deflandre 1937) Stover & Evitt 1978
(Plate 15, fig. 7)
1933a Hystrichosphaera xanthiopyxides Wetzel, p. 44,
45, plate 4, fig. 25 (name not validly
published).
1937 Hystrichosphaeridium xanthiopyxides Wetzel ex
Deflandre, p. 77.
1965 Baltisphaeridium xanthiopyxides (Wetzel ex
Deflandre) – Downie & Sarjeant, p. 98.
1968 Hystrichosphaeridium? xanthiopyxides (Wetzel
ex Deflandre) – Morgenroth, p. 556.
1969 Prolixosphaeridium? xanthiopyxides (Wetzel ex
Deflandre) – Davey et al., p. 17.
1978 Tanyosphaeridium xanthiopyxides (Wetzel ex
Deflandre) – Stover & Evitt, p. 85.
Age. LO: Danian.
Remarks. Fensome et al. (2009, p. 62) is followed here
in using the earliest proposed name, Tanyosphaeridium
xanthiopyxides, for all the Late Cretaceous – Paleocene
forms of Tanyosphaeridium.
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72
Genus Taurodinium gen. nov.
Type. Plate 14, fig. 13, as Taurodinium granulatum.
Etymology. The name derives from the Latin taurus, mean -
ing bull, in reference to the two prominent apical horns.
Description. Dorso-ventrally compressed ceratiacean
cysts with an autocyst having six horns: two apical, two
lateral and two antapical, all with closed, acuminate to
slightly rounded terminations. Autophragm smooth to
finely granulate or perforate and very thin. Parata bula -
tion not expressed. Cingulum may be indicated above
the two lateral horns but there is no indication of a
sulcus. Archaeopyle apical with a straight to weakly
angular margin; operculum usually attached.
Remarks. Nyktericysta and Vesperopsis have the same thin
and fragile appearance as Taurodinium, but have only a
single apical horn. Satyrodinium Lentin & Ma num
1986, has two to three apical and one or more antapical
horns, but is a peridiniacean cyst with an intercalary
archaeopyle, lacks lateral horns, and is cavate. The
acritarch genus Limbicysta Marshall 1989, from Upper
Cretaceous non marine to nearshore environments, has
an elongate split protrusion at one end that shows some
resemblance to the apical horns of Taurodinium but
Limbicysta lacks a regular opening and other protrusions.
Taurodinium granulatum sp. nov.
(Plate 14, figs 13–15, 17, 18)
1992 Gen. et sp. indet. Piasecki et al., fig. 6L, M.
1995 Nyktericysta sp. Gregory & Hart, plate 7, figs 6, 7.
2011 Gen. et sp. indet. Piasecki et al. 1992 – Nøhr-
Hansen et al., figs 3, 4.
2012 Gen. et sp. indet. Piasecki et al. 1992 – Nøhr-
Hansen, plate X, figs 12–19.
Holotype. Plate 14, fig. 13, from a cuttings sample at 2340
m in Ikermiut-1, offshore West Greenland, MGHU no.
31333, sample Ikermiut 2289, slide s-261-3, England
Finder T19-1. Size: overall length 105 μm, length of body
54 μm, body width 35 μm, length of apical horns up to
30 μm, length of lateral horns 13 μm, length of antapical
horns 22 μm, wall less than 1 μm thick. The age deter-
mined for the sample from which the holotype was recov-
ered is late Thanetian.
Etymology. The epithet is from the Greek granulatum
meaning granulate, in reference to the surface ornamen-
tation.
Description. A species of Taurodinium with two apical
horns of approximately equal length, two short lateral
horns and two antapical horns of unequal length with the
left being the longer. The antapical horns narrow at about
one-third along their length. Autophragm smooth to fine-
ly granulate, thin, hyaline and often wrinkled. The apical
horns appear to be oriented in a plane perpendicular to
that of the other horns, giving the cyst a twisted look.
Size. Overall length 75–105 μm, length of body 40–70
μm, body width 28–60 μm, length of apical horns 15–
25 μm, length of lateral horns 7–17 μm, length of
antapical horns 10–32 μm; seven specimens measured.
Age. LO: Bartonian; peak early Ypresian.
Remarks. Taurodinium granulatum was originally record -
ed as ‘Gen. et sp. indet.’ by Piasecki et al. (1992) from a
miospore-dominated palynological assemblage devoid
of other dinocysts. The assemblage was from a silty shale
clast in subaqueous volcanic breccias from the lower
Rinks Dal Member, Maligât Formation, West Green -
land; the basalts were radiometrically dated as 61.2 ± 0.4
Ma, and thus of Selandian age (Larsen et al. 2015).
Gregory & Hart (1995) recorded a specimen of Ny k -
tericysta sp. that may be assignable to Taurodinium gra -
nulatum from sediments dated as Thanetian. Nøhr-
Hansen et al. (2011) recorded Gen. et sp. indet. of
Piasecki et al. (1992) from Thanetian – lower? Ypresian
non-marine or marginal marine deposits of north-eastern
Greenland. Recently, this species has been found to be
common in the Thanetian‒Ypresian non-marine or
marginal marine Kulhøje Member in the Kangerlussuaq
Basin, south-eastern Greenland (Nøhr-Hansen 2012). In
the present study, Taurodinium granulatum was encoun-
tered in wells on the West Greenland continental margin
in samples together with the Late Paleocene marker
Axiodinium augustum, as well as in samples of Ypresian
age from the Saglek Basin. However, the species has been
mostly recorded from spore-dominated or marginal
marine, algae-dominated assemblages. We thus follow
Nøhr-Hansen (2012) in regarding it as a freshwater to
brackish-water indicator.
Genus Tenua Eisenack 1958
Type. Eisenack 1958, plate 23, fig. 1, as Tenua hystrix.
1958 Tenua Eisenack, p. 410.
1981 Cerbia Below, p. 8.
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73
Remarks. The genus is distinguished from Cyclone p he -
lium (incorporating Circulodinium) by having some
processes showing alignment in penitabular rows. Aptea
differs in having three horns – an apical, an antapical,
and a third, probably postcingular – of variable devel-
opment; the probable postcingular horn is always re -
duced. Cerbia is considered here to be a taxonomic
junior synonym of Tenua.
Tenua hystrix Eisenack 1958
(Plate 14, figs 19, 20)
1958 Tenua hystrix Eisenack, p. 410, plate 23, figs
1–4; text-fig. 10.
1972 Tenua hystricella Eisenack & Kjellström,
p. 1039.
1978 Cyclonephelium hystrix (Eisenack) – Davey,
p. 894.
Age. LO: late Aptian.
Remarks. Sarjeant (1985b, p. 94, 95) retained this
species in Tenua. The distribution of processes in this
species is variable. The material in this study includes
forms in which processes are largely absent on mid-
ventral and mid-dorsal surfaces (for example that
shown in Plate 14, fig. 19).
Genus Thalassiphora Eisenack & Gocht 1960
Type. Eisenack 1954, plate 12, fig. 17, as Pterosper m -
opsis pelagica.
1960 Thalassiphora Eisenack & Gocht, p. 513.
1966a Erikania Morgenroth, p. 27.
1980 Subathua Khanna & Singh, p. 307, 308.
Remarks. Fensome et al. (2009, p. 62) provided a com -
prehensive synopsis for Thalassiphora based on the
emendations of the genus by Williams & Downie
(1966a, p. 234), Gocht (1968, p. 153) and Benedek &
Gocht (1981, p. 59).
Thalassiphora delicata Williams & Downie 1966a
(Plate 15, fig. 1)
1966a Thalassiphora delicata Williams & Downie,
p. 235, plate 26, fig. 8.
1973 Disphaeria delicata Norvick, p. 43.
Age. LO: middle Bartonian. Not plotted.
Remarks. Lentin & Williams (1977a, p. 54) retained
this species in Thalassiphora.
Thalassiphora fenestrata Liengjarern et al. 1980
(Plate 15, figs 2–4)
1980 Thalassiphora fenestrata Liengjarern et al.,
p. 489, plate 54, fig. 1.
Age. LO: earliest Rupelian.
Remarks. Thalassiphora fenestrata has large fenestra-
tions, which are restricted to lateral and ventral areas of
the pericyst. Liengjarern et al. (1980, p. 489) consid-
ered its stratigraphic range to be late Eocene to early?
Oligocene.
Thalassiphora pelagica (Eisenack 1954) Eisenack &
Gocht 1960
(Plate 15, figs 5, 6)
1954 Pterospermopsis pelagica Eisenack, p. 71, plate
12, figs 17, 18.
1960 Thalassiphora pelagica (Eisenack) – Eisenack
& Gocht, p. 513, 514.
1966 Thalassiphora sueroi Pöthe de Baldis, p. 224,
225, plate 2, fig. d.
1973 Disphaeria pelagica (Eisenack) – Norvick,
p. 46.
1981 Disphaeria sueroi (Pöthe de Baldis) – Yun
Hyesu, p. 70.
Age. LO: Chattian.
Remarks. Lentin & Williams (1977a, p. 54) retained
this species in Thalassiphora.
Genus Trichodinium Eisenack & Cookson 1960
Type. Eisenack & Cookson 1960, plate 2, fig. 4, as
Trichodinium pellitum.
1960 Trichodinium Eisenack & Cookson, p. 5.
Remarks. Trichodinium is a spheroidal to ovoidal prox-
imate gonyaulacacean cyst that apically can have a horn
or several spines. The autophragm may be tabulate, as
determined from the alignment of some of the nume -
rous short spines or bifid processes. Its surface has been
described as fibro-pitted (see Fensome et al. 2009, p.
64). The archaeopyle is precingular, with the formula
P3´´; the operculum is free. Fensome et al. (2009, p. 64)
Bulletin36.qxp_Bulletin 36 19/12/16 13.40 Side 73
74
discussed the morphological similarity between Tricho -
dinium and Xenicodinium Klement 1960. They recom-
mended that Xenicodinium should be restricted to its
type material and all forms with the appropriate mor -
phology should be included in Trichodinium.
Trichodinium castanea Deflandre 1935 ex Clarke
& Verdier 1967
(Plate 15, fig. 8)
1935 Palaeoperidinium castanea Deflandre, p. 229,
plate 6, fig. 8; name not validly published.
1967 Trichodinium castanea (Deflandre) – Clarke &
Verdier, p. 19, 20.
Age. LO: Campanian.
Remarks. Some tabulation can usually be discerned on
Trichodinium castanea, which is also distinguished by
its dense covering of short, acuminate to bifid spines.
Genus Trithyrodinium Drugg 1967
Type. Drugg 1967, plate 3, fig. 2, as Trithyrodinium evittii.
1967 Trithyrodinium Drugg, p. 20.
Synopsis. Peridiniacean (deflandreoid) cysts that are
proxi mate and rounded to peridinioid in shape, with an
antapex that is rounded, symmetrical or, more usually
weakly to strongly asymmetrical, the left side being
larger; endophragm rounded pentagonal to subcircular
in dorso-ventral outline. Cavate; endocyst sometimes
strongly developed with a fragile or thin-walled peri -
phragm that is easily lost. Archaeopyle intercalary, with
formula I(1–3a); operculum free, compound. Plate 2a is
always iso- to steno deltaform.
Remarks. The synopsis is largely a repeat of that provid-
ed by Fensome et al. (2009, p. 64), though with the
concept broadened to include rounded forms and to
recognise that the pericyst may be durable. The 2a
plate, which is revealed through archaeopyle develop-
ment, can show some variation in width but is always
deltaform. The genus Pierceites also has a 3I archaeopyle
but the endocyst is absent or weakly developed.
Trithyrodinium? conservatum sp. nov.
(Plate 15, figs 9–14)
2003 Deflandrea sp.1 Nøhr-Hansen, plate 3, figs 4–6.
Holotype. Plate 15, fig. 12 and Nøhr-Hansen (2003,
plate 3, fig. 6), from a sidewall-core sample at 1155 m in
Ikermiut 1, MGHU no. 26502, sample 04E006504,
slide 2, co-ordinates 36.0 × 97.1, England Finder U36/1.
Pericyst length 60 μm, width 63 μm, endocyst length 44
μm, width 52 μm. The age determined for the sample
from which the holotype was recovered is Lutetian.
Etymology. The epithet is from the Latin conservatus,
meaning retain or conserve, in reference to the constant
presence of the pericyst.
Description. A species of Trithyrodinium with a com mon -
ly rounded but sometimes ovoidal pericyst that is always
present. The endocyst generally mimics the shape of the
pericyst. As a rule the cyst is circumcavate, but the endo-
cyst may occasionally be in partial contact with the peri-
cyst. The periphragm and endophragm are both thin, at
the most slightly over 1 μm thick. The periphragm varies
from laevigate (the usual condition), to faintly granulate
or verrucate. The I to 3I archaeopyle is formed from the
loss of one to three intercalary plates individually, any of
which can remain attached posteriorly.
Size. Pericyst length 48–60 μm, width 49–65 μm,
endo cyst length 44–51 μm, width 43–56 μm; seven
specimens measured.
Age. LO: Lutetian. Not plotted.
Remarks. Trithyrodinium? conservatum is unusual in that
the pericyst is always preserved. When the pericyst has a
sub dued granulate or verrucate ornamentation, this tends
to be restricted to the mid-dorsal and mid-ventral regions.
Folds are consistently present on the pericyst but appear to
be random. The exact nature of the archaeopyle is unclear.
In some specimens its polygonal shape appears to indicate
that multiple plates are missing, but in others it appears to
reflect loss of a single intercalary plate. Because of the
uncertainty regarding the archaeopyle, the species is only
assigned questionably to Trithyrodinium.
Trithyrodinium evittii Drugg 1967
(Plate 15, figs 17–19)
1967 Trithyrodinium evittii Drugg, p. 20, plate 3,
figs 2, 3; plate 9, fig. 2.
1969b Trithyrodinium fragile Davey, p. 11, plate 3,
figs 6, 9.
Age. LO: Danian.
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75
Trithyrodinium quinqueangulare Marheinecke
1992
(Plate 15, fig. 16)
1992 Trithyrodinium quinqueangulare Marheinecke,
p. 95, plate 19, figs 9–11.
Age. LO: Maastrichtian.
Remarks. Trithyrodinium quinqueangulare has a pericyst
and endocyst, which are sometimes closely appressed.
Both pericyst and endocyst are pentagonal and have two
more or less equal antapical protuberances. The cingu-
lum is clearly delineated. Plate 4´´ is reduced in an ante-
rior–posterior direction. Some of the Labrador Margin
specimens have a verrucate endophragm.
Trithyrodinium suspectum (Manum & Cookson
1964) Davey 1969b
(Plate 15, fig. 15)
1964 Hexagonifera suspectum Manum & Cookson,
p. 9, 10, plate 1, figs 9–13.
1969b Trithyrodinium suspectum (Manum &
Cookson) – Davey, p. 12.
Age. LO: Campanian.
Remarks. Trithyrodinium suspectum has a thick granular
endophragm, which appears to be tectate. The archaeo -
pyle may form from the detachment of the 1a and 3a
plates, whereas the 2a plate can remain attached poste-
riorly, along its boundary with the 4´´ plate. One spe -
cimen of Trithyrodinium suspectum illustrated in Ma-
num & Cook son (1964, plate 1, fig. 11) has a steno -
deltaform 2a plate rather than the more commonly
observed isodeltaform 2a.
Genus Tuberculodinium Wall 1967
Type. Rossignol 1962, plate 2, fig. 1, as Pterospermopsis?
vancampoae.
1967 Tuberculodinium Wall, p. 114.
Synopsis. Goniodomacean (gambierdiscoid) cysts prox-
imate, preicyst and endocyst both discoidal to subsphe -
roidal. Holocavate. Numerous intratabular pillar- to
barrel-shaped processes support an ectophragm.
Archae o pyle antapical, involving usually 2–3 para-
plates, operculum free, compound.
Remarks. Tuberculodinium represents the cysts of Pyro -
phacus, with its multiplate tabulation; the ectophragmal
supports (processes), although numerous, can be readily
interpreted as intratabulate.
Tuberculodinium vancampoae (Rossignol 1962)
Wall 1967
(Plate 15, fig. 20)
1962 Pterospermopsis? vancampoae Rossignol, p.
134, plate 2, fig. 1.
1967 Tuberculodinium vancampoae (Rossignol) –
Wall, p. 114, 115.
1971 Pyrophacus vancampoae (Rossignol) – Wall &
Dale, p. 234.
Age. LO: Tortonian?
Remarks. Head (1996, p. 1232) retained this species in
Tuberculodinium.
Genus Vesperopsis Bint 1986
Type. Bint 1986, plate 5, figs 9, 12–13; text-fig. 5, as
Vesperopsis mayii.
1986 Vesperopsis Bint, p. 156.
Remarks. Vesperopsis is a proximate ceratiacean cyst with
an autophragm that has at least three horns – one apical
and two antapical. Commonly, it may also have equa-
torial horns with or without pre- and postcingular
branches. The archaeopyle is apical, with the formula
A(1–4´); the operculum is usually attached. Vesperopsis
was emended by Qiao Xiuyun et al. (1992, p. 32, 33,
36, 37), Wan Chuanbiao & Qiao Xiuyun (1994, p.
503) and Mao Shaozhi et al. (1999, p. 149, 150).
However, we adhere to the synopsis of Fensome et al.
(2009, p. 65).
Vesperopsis longicornis (Batten & Lister 1988)
Harding 1990
(Plate 16, figs 1, 2)
1988 Australisphaera longicornis Batten & Lister, p.
340, 341, fig. 1b–e, g.
1990 Vesperopsis longicornis (Batten & Lister) –
Harding, p. 21.
Age. LO: Albian.
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76
Remarks. Harding (1990, p. 21) emended the diagnosis of
Vesperopsis longicornis, based on it having an auto phragm
and an attached operculum. According to Har ding, an
unusual aspect of Vesperopsis longicornis is that the opercu-
lum appears to be attached dorsally; operculum attachment
in ceratiacian cysts, where present, is almost always ventral.
Genus Wallodinium Loeblich Jr. & Loeblich III
1968
Type. Cookson & Eisenack 1960b, plate 39, fig. 4, as
Diplotesta glaessneri.
1960b Diplotesta Cookson & Eisenack, p. 256; name
illegitimate.
1968 Wallodinium Loeblich Jr. & Loeblich III, p. 212.
Synopsis. Proximate gonyaulacalean cyst, elongate cres-
cent-shaped to subcylindrical. Bicavate, circumcavate
or epicavate. Pericyst much longer than endocyst; both
bodies generally rounded apically and antapically, al -
though the endocyst may have short horns. Tabulation
indicated by apical archaeopyle, with inferred formula
of A(1–4´); peri- and endoarchaeopyle free or attached.
Sometimes an equatorial constriction or ornamentation
alignment marks a cingulum. Wall smooth or with orna -
mentation of low relief.
Remarks. Wallodinium was considered to be an acritarch
by Duxbury (1983, p. 68) and Fensome et al. (1990, p.
535). However, Riding (1994, p. 17, 18) emended the
diagnosis to include reference to the indications of
gonyaulacalean tabulation and the presence of an apical
archaeopyle; the above synopsis is based on Riding’s
emendation.
Wallodinium luna (Cookson & Eisenack 1960a)
Lentin & Williams 1973
(Plate 16, fig. 5)
1960a Diplotesta luna Cookson & Eisenack, p. 10,
11, plate 3, fig. 21.
1973 Wallodinium luna (Cookson & Eisenack) –
Lentin & Williams, p. 140.
Age. LO: Campanian.
Genus Wetzeliella Eisenack 1938 emend. Williams,
Damassa, Fensome & Guerstein in Fensome et al.
2009
Type. Eisenack 1938, fig. 4, as Wetzeliella articulata.
1938 Wetzeliella Eisenack, p. 187.
1979 Gochtodinium Bujak, p. 310–312.
2009 Wetzeliella Eisenack – emend. Williams,
Damassa, Fensome & Guerstein in Fensome
et al., p. 65.
Synopsis. Wetzelielloidean cysts with a soleiform archae -
o py le and processes that are predominantly non tabular,
al though they can show some alignment, and are dis -
tally free.
Wetzeliella articulata Wetzel in Eisenack 1938
emend. Williams, Damassa, Fensome & Guerstein
in Fensome et al. 2009
(Plate 16, figs 3, 4)
1938 Wetzeliella articulata Wetzel in Eisenack,
p. 187, text fig.4.
1938 Palaeoperidinium articulatum Wetzel in
Eisenack, p. 187 (name not validly
published).
1948 Hystrichosphaeridium articulatum (Wetzel in
Eisenack) – Pastiels, p. 42.
1960 Wetzeliella echinulata Vozzhennikova, plate 3,
fig. 3 (name not validly published).
1967 Rhombodinium coronatum Vozzhennikova,
p. 170, 171, plate 89, figs 1–3, 5; plate 90,
figs 1–5.
1967 Wetzeliella echinulata Vozzhennikova, p. 164,
165.
1975 Wetzeliella horrida Jan du Chêne &
Châteauneuf, p. 28, 30, plate 1, figs 1–7;
plate 3, figs 1–6.
1976 Wetzeliella coronata (Vozzhennikova) – Lentin
& Williams, p. 131.
Age. LO: Rupelian. Not plotted.
Remarks. According to the literature, Wetzeliella artic-
ulata has a stratigraphic range of Eocene–Oligocene.
But when the species is restricted to wetzeleilloideans
with a soleiform archaeopyle, intratabular processes and
five horns, its stratigraphic range is more restricted,
being Bartonian to Rupelian.
Genus Xenascus Cookson & Eisenack 1969
Type. Cookson & Eisenack 1969, fig. 1I, J, as Xenascus
australensis.
1969 Xenascus Cookson & Eisenack, p. 7.
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77
Remarks. Fensome et al. (2009, p. 67) provided a com -
prehensive synopsis of Xenascus in which they referred
to lateral rather than postcingular, horn(s) because they
seem to emanate from the cingulum as well as postcin-
gular areas. Fensome et al. (2009) also discussed the
emendations of Yun Hyesu (1981, p. 60) and Stover &
Helby (1987, p. 128).
Xenascus ceratioides (Deflandre 1937) Lentin
&Williams 1973
(Plate 16, figs 9, 10)
1937 Hystrichosphaera ceratioides Deflandre, p. 66,
67, plate 12 (also labelled as plate 9), figs 7, 8.
1967 Pseudoceratium ceratioides (Deflandre) –
Clarke & Verdier, p. 60.
1970 Spiniferites ceratioides (Deflandre) – Sarjeant,
p. 76.
1971 Phoberocysta ceratioides (Deflandre) – Davey
& Verdier, p. 26.
Age. LO: Campanian.
Remarks. Xenascus ceratioides has three horns: one api -
cal, one lateral and one antapical. The horns are not
perforated.
Genus Fromea (Cookson & Eisenack 1958) Yun
Hyesu 1981
Type. Cookson & Eisenack 1958, plate 5, fig. 10, as
Fromea amphora.
1958 Fromea Cookson & Eisenack, p. 55.
1973 Xenascus ceratioides (Deflandre) – Lentin &
Williams, p. 144.
Remarks. Although many workers assign Fromea to the
dinoflagellates, some (e.g. Duxbury 1980, p. 134; Fen -
some et al. 1990, p. 227; Fensome & Williams 2004, p.
742) consider it to lack unequivocal morphological
indications of dinoflagellate affinity and thus prefer to
consider it an acritarch.
Xenascus wetzelii Slimani 1996 ex Slimani 2001a
(Plate 16, figs 6–8)
1985 Odontochitina wetzelii Wilson in Foucher in
Robaszynski et al., p. 33, plate 10, figs 9–12;
name not validly published.
1996 Xenascus wetzelii Slimani, p. 380, 381, plate
3, figs F, G; plate 4, figs A, B; text-fig. 7A, B;
name not validly published.
2001a Xenascus wetzelii Slimani 1996 ex Slimani,
p. 9, plate 2, figs 3, 4.
Age. LO: Campanian.
Remarks. Xenascus wetzelii is cornucavate to circumca-
vate. It has a long postcingular horn and a long antapi-
cal horn, both of which are perforate distally and can be
acuminate or bifurcate at their extremities. Tabulation
is clearly shown by parasutural crests and gonal process-
es, the latter being acuminate, bifurcate or trifurcate.
Slimani (2001a, p. 9; 2001b, p. 194) considered Odon -
tochitina wetzelii to be a taxonomic junior synonym of
Xenascus wetzelii.
Fromea nicosia Jansonius 1989
(Plate 16, fig. 19)
1989 Fromea nicosia Jansonius, p. 67, plate 1,
figs 2–7; text-fig. 1.
Age. LO: early Campanian.
Fromea quadrangularis sp. nov.
(Plate 16, figs 11, 12, 15, 16)
Holotype. Plate 16, fig. 16, from a cuttings sample at
2375 m in Skolp E-07, MGUH no. 31352, sample
YD15665, slide 04, co-ordinates 30.9 × 111.9, Eng land
Finder D31-1. Overall length 88 μm; maximum width
35 μm; minimum width 27 μm; wall thickness 1 μm.
Systematics – acritarchs and other algae
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The age determined for the sample from which the holo-
type was recovered is early Campanian, Late Cretaceous.
Etymology. The epithet is from the Latin words quadra,
meaning four and angularis, having angles, in reference
to the quadrangular shape of specimens of this species.
Diagnosis. A slender elongate rectangular species of
Fro mea with a thin smooth wall and generally two
longitudinal folds. Where present, the folds start at the
antapex and continue along the body to the circular
‘apical’ opening. No accessory opening sutures or equa-
torial ‘girdle’ have been observed.
Size. Overall length 77–101 μm; maximum width 30–
45 μm; minimum width 27–37 μm; seven specimens
measured.
Age. LO: early Campanian.
Remarks. Some specimens of Fromea quadrangularis are
twisted, resulting in an elongate jar-like shape with
longitudinal folds; other specimens have been slightly
compressed, resulting in a weak equatorial extension
and a splitting of the elongate folds.
Genus Microsphaeridium Benedek 1972
Type. Benedek 1972, plate 12, fig. 3a, b, as Micro -
sphaeridium ancistroides.
1972 Microsphaeridium Benedek, p. 46, 47.
Remarks. Benedek & Sarjeant (1981, p. 346, 347)
interpreted Microsphaeridium ancistroides as “the de -
tached opercula of skolochorate dinoflagellate cysts.”
Pending further investigation, we prefer to consider
Microsphaeridium to be an acritarch.
Microsphaeridium ancistroides Benedek 1972
(Plate 16, figs 13, 14, 17, 18)
1972 Microsphaeridium ancistroides Benedek, p. 47,
plate 12, fig. 3a, b; text-fig. 21.
Age. LO: middle Miocene or younger; not well
constrained. Not plotted.
Genus Palambages Wetzel 1961
Type. Wetzel 1961, plate 1, fig. 11, as Palambages mo -
ru losa.
1961 Palambages Wetzel, p. 338.
Remarks. Wetzel (1961) noted that the microfossils as -
signed to Palambages are identical to forms recorded as
“Morulosae” in Wetzel (1933a, p. 23, 24, plate 4, figs
1–5). Wetzel (1933a) had compared these fossils with
certain colonial algae, but noted their likeness to the
egg-balls of planktonic crustaceans. Manum & Cook -
son (1964, p. 23) considered them to represent colonies
of green algae.
Palambages spp.
(Plate 16, fig. 20)
Age. Range not determined. Not plotted.
Genus Paralecaniella Cookson & Eisenack 1970a
Type. Deflandre & Cookson 1955, plate 9, fig. 6, as
Epicephalopyxis indentata.
1970a Paralecaniella Cookson & Eisenack, p. 323.
Remarks. Understanding of this genus, which is abun-
dant in what seem to be shallow-water environments, is
tenuous. Elsik (1977, p. 96), who emended the diagno-
sis of the genus and the type, considered Paralecaniella
to be a dinocyst. We agree with Fensome et al. (2009,
p. 67) that it is probably an algal cyst of non-dinoflagel-
late affinity.
Paralecaniella indentata (Deflandre & Cookson
1955) Cookson & Eisenack 1970a
(Plate 17, figs 1, 2)
1955 Epicepholopyxis indentata Deflandre &
Cookson, p. 292, plate 9, figs 5–7; text-fig. 56.
1970a Paralecaniella indentata (Deflandre &
Cookson) – Cookson & Eisenack, p. 323.
1973 Scriniodinium? nilsii Kjellström, p. 42, fig. 35.
Age. Peak occurrence within the earliest Ypresian.
Remarks. The peak occurrence of Paralecaniella inden-
tata is close to the Paleocene–Eocene boundary.
Genus Pediastrum Meyen 1829
Type. Meyen 1829, plate 43, figs 6–20, as Pediastrum
duplex.
1829 Pediastrum Meyen, p. 772.
Remarks. Pediastrum is a nonmotile coenobial green
alga that is found in freshwater environments.
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79
Pediastrum spp.
(Plate 17, figs 3, 4)
Age. Range not determined. Not plotted. The range of
the genus extends from Early Cretaceous to Recent
(Batten 1996).
Remarks. The presence of Pediastrum spp. in some of the
samples from marine deposits in the offshore wells is
indicative of offshore transport of freshwater elements.
Genus Tetraporina Naumova 1939
Type. Tetraporina antiqua Naumova 1950, designated
by Potonié (1960, p. 130).
1939 Tetraporina Naumova, p. 357.
1956 Tetrapidites Klaus in Meyer, p. 107.
1960 Tetraporopollenites Frantz, p. 559.
1963 Balmeella Pant & Mehra, p. 116.
1980 Tetraporina Naumova – emend. Lindgren,
p. 346.
Synopsis. Acid resistant, unicellular, tetrahedral or par -
al lel epipedal microfossils with or without obvious pore
or other dehiscence mechanism. Wall single or double
layered.
Remarks. Described originally as representing pollen,
Hemer & Nygreen (1967) considered Tetraporina to be
an algal genus, and it has been considered generally to
represent the acritarchs or algae since then. The above
synopsis largely follows the emended diagnosis of
Lindgren (1980). Recognition of Tetaporina is based
mainly on its distinctive tetrahedral or parallelepipedal
shape (a parallelepiped is a geometric figure with six
faces, all parallelograms and all opposite faces being simi-
lar and parallel). Although most commonly related to
modern zygospores of the Zygnemataceae, Lindgren
(1980) considered Teraporina to be polyphyletic and
difficult to match with particular modern algae because
of its simple shape, rigours of preservation and ontoge-
netic factors. Most algae to which Tetraporina can be
related are freshwater. The stratigraphic range of the
genus extends from Carboniferous to Quaternary.
There has been considerable debate about the
nomenclatural status of Tetraporina. Jansonius & Hills
(1981, card 3917) considered that the name was not
validly published in Naumova (1939) because it was
proposed in anticipation of future acceptance of the
name (see McNeill et al. 2012, Article 36.2). However,
the case for this has not proved compelling for other
authors (Lindgren 1980; Farr & Zijlstra 1996).
Although both sets of arguments have merit, for prag-
matic reasons, here we accept Tetraporina as valid in
Naumova (1939).
Tetraporina sp. A
(Plate 17, figs 5–7)
Description. A form of Tetraporina with a tetrahedral
outline and indentations at the apices, although these
are not clearly perforated by pores. The apices have no,
or only short, extensions. The wall surface is smooth or
has subdued ornament such as small verrucae. This
form may correspond with one or more of the species
described by Lindgren (1980), but morphological over-
lap, preservation and flexibility of the wall, and the
quality of Lindgren’s illustrations make comparison
difficult.
Age. LO: earliest Rupelian.
Tetraporina? sp. B
(Plate 17, figs 8–12)
Description. A form with broad truncated extensions at
each of the four apices. The extensions may be relatively
short and converge on a central rhombic area, or they
may just converge centrally without the development of
a central rhombic area, in which case the entire speci-
men is essentially a cross-like structure. Tetraporina
does not typically have extensions, so this form is only
tentatively associated with that genus.
Age. Not plotted.
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Miospores
Genus Afropollis Doyle et al. 1982
Type. Brenner 1968, plate 10, fig. 5, as Reticulato -
sporites jardinus.
1982 Afropollis Doyle et al., p. 44.
Afropollis sp.
(Plate 17, figs 18, 19)
Remarks. This form is assigned to Afropollis because of
its possession of a network of reticulate to regulate muri
separated from the nexine. The grains are apparently
inaperturate. No ring furrow was discerned. The net -
work of muri is considerably coarser than that exhibited
by Afropollis jardinus, Afropollis operculatus and Afro -
pollis zonatus.
Age. LO: Cenomanian.
Genus Appendicisporites Weyland & Krieger 1953
Type. Weyland & Krieger 1953, plate 11, fig. 54, as
Appendicisporites tricuspidatus.
1953 Appendicisporites Weyland & Krieger, p. 12.
Appendicisporites potomacensis Brenner 1963
(Plate 17, fig. 20)
1963 Appendicisporites potomacensis Brenner, p. 46,
plate 6, figs 4, 5.
1985 Plicatella potomacensis (Brenner) – Davies,
p. A49.
Age. LO: mid-Cretaceous, but not well constrained.
Not plotted.
Remarks. This species is retained in Appendicisporites,
following Burden & Hills (1989, p. 100) and Nichols
& Sweet (1993, p. 548).
Appendicisporites unicus (Markova in Ivanova &
Markova 1961) Singh 1964
(Plate 18, fig. 1)
1961 Anemia unica Markova in Ivanova &
Markova, p. 53, plate 20, fig. 3a, b.
1964 Appendicisporites unicus (Markova in Ivanova
& Markova) – Singh, p. 53.
1985 Plicatella unica (Markova in Ivanova &
Markova) – Davies, p. A 53.
Age. LO: mid-Cretaceous, but not well constrained.
Not plotted.
Remarks. This species is retained in Appendicisporites,
following Burden & Hills (1989, p. 100).
Genus Aquilapollenites Rouse 1957 emend.
Braman 2013
Type. Radforth & Rouse 1954, plate 1, fig. 14, as “N2”.
Holotype lost; lectotype (possibly the holotype) select-
ed from restored type slide by Tschudy & Leopold
(1971, plate 2, fig. 1); neotype designated by Srivastava
& Rouse (1970, plate 1, figs 4–7); all as Aquilapollenites
quadrilobus.
1957 Aquilapollenites Rouse, p. 370.
1970 Hemicorpus Krutzsch, p. 107.
2013 Aquilapollenites Rouse – emend. Braman, p.14.
Remarks. Braman (2013) is followed here in consider-
ing Aquilapollenites to comprise triprojectate pollen
that are heteropolar and have isolated sculptural ele -
ments. Other triprojectate pollen found in this study
are assignable to Parviprojectus and Translucentipollis.
Aquilapollenites quadrilobus Rouse 1957
(Plate 17, figs 14, 15, 17)
1957 Aquilapollenites quadrilobus Rouse, p. 371,
plate 2, figs 8, 9.
1961 Aquilapollenites polaris Funkhouser, p. 198,
plate 1, figs 1, 2.
1961 Aquilapollenites pulcher Funkhouser, p. 198,
plate 1, fig. 7a–c.
1970 Mancicorpus polaris (Funkhouser) – Stanley,
p. 30.
1970 Hemicorpus polaris (Funkhouser) – Krutzsch,
p. 107.
1970 Hemicorpus pulcher (Funkhouser) – Krutzsch,
p. 107.
1970 Mancicorpus pulcher (Funkhouser) –
Srivastava, p. 697.
Systematics – miospores and fungal elements
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1970 Aquilapollenites quadrilobus Rouse – emend.
Srivastava & Rouse, p. 1597.
Remarks. The synonymy above follows that in Braman
(2013, p. 28). This widely recorded species is charac-
terised by heteropolar grains with prominent spinate
sculpture. According to Braman (2013) and A. Sweet
(personal communication 2015), this species has a Late
Cretaceous range, so the specimens recorded from the
Labrador–Baffin Seaway are assumed to be reworked.
Age. LO: middle? Eocene (reworked?).
Genus Azolla Lamarck in Lamarck et al. 1783
Type. Azolla filiculoides Lamarck et al. 1783.
1783 Azolla Lamarck in Lamarck et al., p. 343.
Azolla spp.
(Plate 18, figs 2–4)
Age. LO: ?Bartonian. Frequent in late Ypresian.
Remarks. Azolla is a small moss-like, free-floating fresh-
water fern that occurs today in warm climates; it is
famous for its nitrogen-fixing capability. Given the
right combination of time and temperature, sediment
rich in Azolla could represent major sources of oil.
Azolla had coeval blooms in some Arctic and northern
temperate areas during the Early to earliest Middle
Eocene (e.g. Barke et al. 2011, 2012). Barke et al.
(2012) concluded that the presence of the blooms
accords with high-precipitation conditions modelled
for the Early Eocene and implies the presence of exten-
sive wetlands bordering the regional landmasses. They
further suggested that Azolla blooms in the Arctic and
Norwegian Sea basins may indicate widespread fresh
ocean surface waters due to unprecedented discharge of
water from the land and the semi-enclosed nature of
those two basins. The occurrences of Azolla in Labrador
Sea – Davis Strait wells are mostly restricted to a narrow
time interval at the top of the Ypresian. Azolla has not
been recorded in the offshore West Greenland wells.
Evidence of extensive blooms of Azolla in the Lab -
rador–Baffin Seaway was not encountered in this study,
perhaps indicating that this marine basin was broadly
open to the south and had limited or no connection to
the Arctic basin (see Barke et al. 2012, fig. 4 and Nøhr-
Hansen et al. 2016).
The poor preservation and limited frequency of
Azolla specimens in our study have precluded specific
assigments, but several species are probably represented
(see Barke et al. 2012). Previous studies suggest an LO
of Azolla species in the earliest Lutetian.
Genus Baculatisporites Pflug & Thomson in
Thomson & Pflug 1953
Type. Wolff 1934, plate 5, fig. 8, as Sporites priarius.
1953 Baculatisporites Pflug & Thomson in
Thomson & Pflug.
Synopsis. Trilete spores, with a subcircular ambitus; the
laesurae are of variable length, sometimes reaching to
the equator. The ornamentation of the exine is predom-
inantly of bacula, although other types of element may
be present. Distally, the bacula can be flat to multi-
crowned.
Remarks. In our synopsis for Baculatisporites, we allow
for variation in both the width relative to the height of
the baculae, so that the width may exceed the height,
and in the nature of their distal terminations.
Baculatisporites crenulatus sp. nov.
(Plate 18, figs 6–8)
Holotype. Plate 18, fig. 8, from a cuttings sample at
1790–1800 m in Roberval K-92, GSC type collection
no. 137889, sample P17684, slide 01, co-ordinates
10.6 × 104.8, England Finder L34/2. Maximum overall
diameter 65 μm. The age determined for the sample
from which the holotype was recovered is Bartonian.
Etymology. The epithet is from the Latin crenulatus,
meaning minutely crenulate, in reference to the
toothed distal margin of the bacula.
Diagnosis. A species of Baculatisporites with sculptural
elements that have variable distal terminations, ranging
from blunt to rounded to crenulate. There is consider-
able variation in the nature of the distal terminations,
and also in the length-to-width ratio of the elements:
many are of greater length than width and hence true
bacula, but others are of greater width than length.
Size. Diameter 52–65 μm. Length of laesurae 15–18
μm. Bacula height 3–12 μm, width 3–7 μm. Wall
thickness 2–3 μm.
Age. LO: Bartonian. Not plotted.
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Remarks. The ratio of the width to height of the bacu-
lae and their distal variability from blunt to crenulated,
distinguishes Baculatisporites crenulatus from other spe -
cies of the genus.
Genus Callialasporites Dev 1961
Type. Balme 1957, plate 8, fig. 91, as Zonalapollenites
trilobatus.
1961 Callialasporites Dev, p. 48.
1961 Applanopsis Döring, p. 112.
1961 Triangulopsis Döring, p. 113.
1962 Pflugipollenites Pocock, p. 72.
1964 Applanopsipollenites Levet-Carette, p. 107.
1970 Singhiapollis Kar & Sah, p. 107.
Synopsis. Cavate, proximo-distally compressed mio -
spores with a roughly circular to triangular or trilobate
amb. The two wall layers are mostly appressed proxi-
mally and distally, but ambitally and sub-ambitally the
wall layers are variously separated to produce a hollow
zona that may be continuous or constricted to few
(commonly three) to multiple vesicles and/or be radial-
ly plicated. The central body may be circular, triangu-
lar, or irregularly shaped. A proximal, non-functional
triradiate mark may be present.
Remarks. The above description is condensed from a
version provided by Fensome (1983), who gave an
extensive review of this genus and its synonymy.
Callialasporites dampieri (Balme 1957) Dev 1961
(Plate 18, fig. 16)
1957 Zonalapollenites dampieri Balme, p. 32, plate
8, figs 88–90.
1961 Callialasporites dampieri (Balme) – Dev, p.
48.
1961 Applanopsis dampieri (Balme) – Döring, p.
113.
1962 Pflugipollenites dampieri (Balme) – Pocock, p.
72.
1963 Tsugaepollenites dampieri (Balme) –
Dettmann, p. 100.
Remarks. Singh (1971, p. 175) retained this species in
Callialasporites.
Age. LO: Aptian.
Callialasporites obrutus Norris 1969
1969 Callialasporites obrutus Norris, p. 597, plate
110, figs 6, 7.
Age. LO: Aptian.
Genus Caryapollenites Raatz 1938 ex Potonié 1960
Type. Potonié 1931, fig. 2, as Pollenites simplex.
1934 Caryae?-pollenites Potonié & Venitz, p. 21;
name not validly published.
1938 Caryapollenites Raatz, p. 19; name not validly
published.
1960 Caryapollenites Potonié, p. 123.
Remarks. As Nichols & Ott (1978) noted, the nomen-
clatural history of fossil pollen similar to the modern
genus Carya is difficult to unravel. This is also true of
species within Caryapollenites. Nichols & Ott (1978)
developed a biostratigraphic scheme for species of
Caryapollenites and the closely related Momipites for
Paleocene strata of the Wind River Basin of Wyoming.
However, we have found species of these two genera
widely distributed in Paleocene to Miocene sediments,
and our observations do not conform to the tight
Paleocene-restricted ranges proposed by Nichols & Ott
(1978). In our experience, firm ranges for individual
species of these two genera have yet to be determined,
though in the Labrador–Baffin Seaway, Caryapollenites
generally has an LO in the latest Serravallian.
Caryapollenites inelegans Nichols & Ott 1978
(Plate 18, figs 9, 10)
1978 Caryapollenites inelegans Nichols & Ott,
p. 105, 106, plate 2, figs 7, 8.
Age. See ‘Remarks’ under the generic entry for Cary a -
pollenites. Not plotted.
Caryapollenites veripites (Wilson & Webster 1946)
Nichols & Ott 1978
(Plate 18, figs 11, 12)
1946 Carya veripites Wilson & Webster, p. 276, fig. 14.
1978 Caryapollenites veripites (Wilson & Webster) –
Nichols & Ott, p. 106.
Age. See ‘Remarks’ under the generic entry for Carya -
pollenites. Not plotted.
Bulletin36.qxp_Bulletin 36 19/12/16 13.40 Side 82
83
Genus Cerebropollenites Nilsson 1958
Type. Couper 1958, plate 30, fig. 8, as Tsugaepollenites
mesozoicus.
1958 Cerebropollenites Nilsson, p. 72.
Cerebropollenites mesozoicus (Couper 1958)
Nilsson 1958
1958 Tsugaepollenites mesozoicus Couper, p. 155,
plate 30, figs 8–10.
1958 Cerebropollenites mesozoicus (Couper) –
Nilsson, p. 72.
Age. LO: Aptian.
Genus Chenopodipollis Krutzsch 1966
Type. Weyland & Pflug 1957, plate 22, figs 18, 19, as
Periporopollenites multiplex.
1966 Chenopodipollis Krutzsch, p. 35.
Chenopodipollis sp.
(Plate 18, fig. 17)
Age. LO: Miocene?
Genus Cicatricosisporites Potonié & Gelletich 1933
Type. Potonié & Gelletich 1933, plate 1, fig. 1, as
Cicatricosisporites dorogensis, designated by Potonié
(1956, p. 47).
1933 Cicatricosisporites Potonié & Gelletich, p. 522.
1950 Mohrioidites Thiergart, p. 84 (name not valid-
ly published).
1951 Mohrioisporites Potonié, p. 144.
Remarks. Emendations for Cicatricosisporites have been
proposed by Potonié (1966, p. 58) and Dettmann &
Clifford (1992, p. 289–291).
Cicatricosisporites minutaestriatus (Bolkhovitina
1961) Pocock 1964
1961 Pelletieria minutaestriata Bolkhovitina, p. 68,
plate 20, fig. 1a–f; plate 21, fig. 3a–g.
1964 Cicatricosisporites minutaestriatus
(Bolkhovitina) – Pocock, p. 159.
1971 Cicatricosisporites augustus Singh, p. 68, plate
7, figs 3–11; text-fig. 7M.
Age. LO: early Turonian.
Cicatricosisporites ornatus Srivastava 1972
(Plate 19, figs 1–4)
1972 Cicatricosisporites ornatus Srivastava, p. 9, plate
5, figs 3‒11; plate 6, figs 1‒4.
Age. LO: Priabonian.
Remarks. The forms studied here are very similar to the
type material, described by Srivastava (1972) from the
Maastrichtian of Alberta.
Genus Cicatricososporites Pflug & Thomson in
Thomson & Pflug 1953
Type. Selling 1944, plate 4, fig. 44, as Schizaea? eocenica.
1953 Cicatricososporites Thomson & Pflug, p. 61.
1959 Schizaeoisporites Krutzsch, p. 226.
Remarks. Jansonius & Hills (1976) did not agree with
Davies (1985) that Cicatricososporites was a junior
homo nym of Cicatricosisporites and retained the former
as a separate genus.
Cicatricososporites eocenicus (Selling 1944)
Jansonius & Hills 1976
(Plate 18, fig. 5)
1944 Schizaea? eocenica Selling, p. 66, plate 4, fig. 44.
1950 Sporites pseudodorogensis Potonié – Thiergart,
p. 84; name not validly published.
1951 Schizaeolsporites pseudodorogensis (Potonié) –
Potonié, p. 144, plate 20, fig. 19; generic
name not validly proposed.
1953 Cicatricososporites pseudodorogensis (Potonié) –
Thomson & Pflug, p. 61.
1976 Cicatricososporites eocenicus (Selling) –
Jansonius & Hills, card 468.
Age. LO: middle Bartonian.
Remarks. Burden & Hills (1989) recorded Cicatricoso -
sporites eocenicus from the Early Cretaceous, but the
species is most common in the early Cenozoic and espe-
cially the Eocene.
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84
Genus Compositoipollenites Potonié 1951 ex
Potonié 1960
Type. Potonié 1934, plate 5, fig. 25, as Pollenites rhi -
zophorus.
1951 Compositoipollenites Potonié, p. 138; name
not validly published.
1960 Compositoipollenites Potonié, p. 105.
Compositoipollenites sp. B of Williams & Brideaux
1975
(Plate 18, fig. 18)
1975 Compositoipollenites sp. B Williams &
Brideaux, plate 43, fig. 15.
Age. LO: Gelasian.
Remarks. Compositoipollenites sp. B of Williams &
Brideaux (1975) is very similar to the pollen of the
extant genus Ambrosia, being tricolporate and orna-
mented with short (up to about 2 μm) conate spines
that are distally pointed.
Genus Corsinipollenites Nakoman 1965
Type. Thiergart 1940, plate 7, fig. 1 as Pollenites oculus
noctis.
Corsinipollenites oculusnoctis (Thiergart 1940)
Nakoman 1965
(Plate 18, figs 19, 20)
1940 Pollenites oculus noctis Thiergart, p. 47.
1965 Corsinipollenites oculusnoctis (Thiergart) –
Nakoman, p. 156.
Age. LO: Bartonian.
Remarks. Fossil pollen with this morphology have also
been described under the modern plant name Jusseia.
Genus Extratriporopollenites Pflug in Thomson &
Pflug 1952 ex Pflug in Thomson & Pflug 1953
Type. Pflug in Thomson & Pflug 1953, plate 6, fig. 2,
as Extratriporopollenites fractus.
1952 Extratriporopollenites Pflug in Thomson &
Pflug, p. 14, 16; name not validly published.
1953 Extratriporopollenites Pflug in Thomson &
Pflug 1952 ex Pflug in Thomson & Pflug,
p. 69.
Extratriporopollenites spp.
(Plate 19, figs 5–9)
Age. LO: Bartonian.
Genus Graminidites Cookson 1947 ex Potonié
1960
Type. Cookson 1947, plate 15, fig. 41, as Monoporites
(Graminidites) media.
1947 Graminidites Cookson, p. 134; name not
validly published.
1960 Graminidites Cookson ex Potonié, p. 111.
Graminidites sp. A. of Williams & Brideaux 1975
(Plate 19, fig. 12)
1975 Graminidites spp. Williams & Brideaux, plate
47, figs 9, 10.
Age. LO: latest Gelasian.
Genus Momipites Wodehouse 1933
Type. Momipites coryloides Wodehouse 1933, fig. 43.
1933 Momipites Wodehouse, p. 511.
Remarks. See ‘Remarks’ under the generic entry for
Caryapollenites.
Momipites annellus Nichols & Ott 1978
(Plate 18, figs 13, 14)
1978 Momipites annelus Nichols & Ott, p. 103,
plate 1, figs 22–25.
Age. See ‘Remarks’ under the generic entry for Cary a -
pollenites. Not plotted.
Momipites coryloides Wodehouse 1933
(Plate 18, fig. 15)
1933 Momipites coryloides Wodehouse, p. 511, fig. 43.
Age. See ‘Remarks’ under the generic entry for Cary a -
pollenites. Not plotted.
Genus Osmundacidites Couper 1953
Type. Couper 1953, plate 1, fig. 5, as Osmundacidites
wellmanii.
Bulletin36.qxp_Bulletin 36 19/12/16 13.40 Side 84
85
1953 Osmundacidites Couper, p. 20.
Remarks. Osmundacidites is characterised by having a
granulate to finely verrucate exine that is commonly
irregularly organised.
Osmundacidites wellmannii Couper 1953
(Plate 20, fig. 1)
1953 Osmundacidites wellmannii Couper, p. 20,
plate 1, fig. 5.
1959 Baculatisporites wellmannii (Couper) –
Krutzsch, p. 142.
1964 Osmundacidisporites wellmannii (Couper) –
Levet-Carette, p. 98.
1968 Todisporites granulatus Tralau, p. 67, plate 7,
fig. 1.
1972 Osmundacidites araucanus Volkheimer,
p. 120, plate 6, figs 47‒49.
1986 Osmunda sp. in Williams, p. 83, plate 1, fig. 2.
Age. LO: Burdigalian.
Remarks. This species has a long range, starting in the
Mesozoic, but its LO appears to be useful in the
Labrador–Baffin Seaway region. We have established a
Burdigalian LO; Williams (1986) considered that
Osmunda sp. (= Osmundacidites wellmannii) has an LO
in the middle to late Miocene. For a full synonymy list-
ing of Osmundacidites wellmannii, see Fensome (1983,
p. 323, 324).
Genus Parviprojectus Mtchedlishvili in
Samoilovitch & Mtchedlishvili 1961 emend.
Braman 2013
Type. Samoilovitch & Mtchedlishvili 1961, plate 73,
fig. 2, as Parviprojectus reticulatus.
1961 Parviprojectus Mtchedlishvili in Samoilovitch
& Mtchedlishvili, p. 225.
2013 Parviprojectus Mtchedlishvili in Samoilovitch
& Mtchedlishvili – emend. Braman, p. 129.
Remarks. Following Braman (2013), Parviprojectus is
considered to comprise triprojectate pollen that are
isopolar and have a reticulate wall.
Parviprojectus reticulatus Mtchedlishvili in
Samoilovitch & Mtchedlishvili 1961
(Plate 17, fig. 16)
1961 Parviprojectus reticulatus Mtchedlishvili in
Samoilovitch & Mtchedlishvili, p. 226, 227,
plate 73, figs 2, 3.
1961 Aquilapollenites reticulatus Stanley, p. 348,
349, plate 8, figs 1–12.
1970 Integricorpus reticulatus (Mtchedlishvili) –
Stanley, p. 29.
1970 Aquilapollenites (Parviprojectus) reticulatus
(Mtchedlishvili) – Kedves & Király, p. 67.
Remarks. This species is characterised by its fine reticu-
late ornament. The above synonymy follows that of
Bra man (2013, p. 148), with only nomenclaturally
significant entries included.
Age. LO: not determined; not plotted. The illustrated
specimen is from late Paleocene strata in Rut H-11.
According to Braman (2013), this species ranges from
the Campanian to early Paleocene, with a ‘provisional’
occurrence in the late Eocene. The latter may be re -
worked, as could be the present occurrence.
Genus Parvisaccites Couper 1958
Type: Couper 1958, plate 29, figs 5, 6, as Parvisaccites
radiatus.
1958 Parvisaccites Couper, p. 154.
Parvisaccites amplus Brenner 1963
1963 Parvisaccites amplus Brenner, p. 78, 79, plate
28, fig. 1a, b; plate 29, fig. 1a, b.
Age. LO: Aptian.
Parvisaccites radiatus Couper 1958
1958 Parvisaccites radiatus Couper, p. 154, plate
29, figs 5–8; plate 30, figs 1, 2.
Age. LO: Aptian.
Genus Periporopollenites Pflug & Thomson in
Thomson & Pflug 1953
Type. Potonié 1931, plate 2, fig. 1, as Pollenites stigmosus.
1953 Periporopollenites Pflug & Thomson in
Thomson & Pflug, p. 111.
1960 Liquidambarpollenites Raatz 1937 ex Potonié,
p. 134.
Bulletin36.qxp_Bulletin 36 19/12/16 13.40 Side 85
86
Periporopollenites sp.
(Plate 19, figs 10, 11)
Age. LO: early? Miocene. Not plotted.
Genus Pistillipollenites Rouse 1962
Type. Rouse 1962, plate 1, figs 10, 12, as Pistilli polle n -
ites macgregorii.
1962 Pistillipollenites Rouse, p. 206.
Remarks. The distinctive characteristic of this triporate
pollen grain is the pistil-like or bulbose ornamentation
that covers the exine.
Pistillipollenites macgregorii Rouse 1962
(Plate 19, figs 13, 14)
1962 Pistillipollenites macgregorii Rouse, p. 206,
plate 1, figs 8–12.
Age. LO: Bartonian.
Genus Quercoidites Potonié et al. 1950 ex Potonié
1960
Type. Potonié 1931, plate 2, fig. 19, as Pollenites henrici.
1950 Quercoidites Potonié et al., p. 54; name not
validly published.
1960 Quercoidites Potonié et al. ex Potonié, p. 92.
Remarks. The exine of this ovoidal, tricolpate pollen
grain is usually granulate to scabrate. The colpi reach
almost to the poles.
Quercoidites sp.
(Plate 19, figs 15, 16)
1986 Quercus form A of Williams, p. 83, plate 1, fig. 8.
Age. LO: Frequent in latest Messinian.
Remarks. This is a finely granular form of Quercoidites,
also found by Williams (1986) and called Quercus form
A. Williams found that the LO of this form was in her
Fagus granulata Zone, which she dated as early
Miocene. Quercoidites sp. differs from Quercoidites sp.
A of Williams & Brideaux (1975) (equivalent to Quer -
c us form B of Williams 1986) in its much finer orna-
ment.
Genus Rugubivesiculites Pierce 1961
Type. Pierce 1961, plate 2, fig. 57, as Rugubivesiculites
convolutes.
1961 Rugubivesiculites Pierce, p. 39.
Remarks. Rugubivesiculites is a bivesiculate pollen with
a distinctive rugulate ornamentation on the proximal
surface of the central body.
Rugubivesiculites spp.
(Plate 20, fig. 2)
Age. LO: Turonian.
Remarks. No attempt was made to speciate the speci-
mens of Rugubivesiculites seen in the Labrador Margin
and offshore West Greenland samples.
Genus Tiliaepollenites Potonié 1931
Type. Potonié 1931, fig. 14, as Tiliaepollenites indu -
pitabilis.
1931 Tiliaepollenites Potonié, p. 4.
1938 Tiliaepollenites Raatz, p. 27.
1953 Intratriporopollenites Pflug & Thomson in
Thomson & Pflug, p. 87.
Tiliaepollenites crassipites (Wodehouse 1933)
comb. nov.
(Plate 19, figs 17, 18)
1933 Tilia crassipites Wodehouse, p. 515, fig. 48.
1969 Tilliaepollenites crassipites (Wodehouse) –
Penny, p. 355 (combination not validly
published, basionym not fully referenced).
1975 Bombacacidites sp. A Williams & Brideaux,
plate 46, fig. 10.
1986 Tilia crassipites Williams, plate 2, fig. 1.
Age. LO: Serravallian.
Remarks. A search of the Palynodata database (Palyno -
data Inc. & White 2008) provided no indication of a
previous formal transfer of this species to Tiliaepollenites.
Tiliaepollenites sp. A
(Plate 19, figs 19, 20)
Description. This form is distinctive in its possession of
a pad-like thickening at the inner end of each colpus.
Bulletin36.qxp_Bulletin 36 19/12/16 13.40 Side 86
87
Age. LO: early? Miocene (not well constrained). Not
plotted.
Genus Translucentipollis Khlonova 1961
Type. Khlonova 1961, plate 16, fig. 121, as Aquila -
pollenites plicatilis.
1961 Translucentipollis Khlonova, p. 89.
1966 Translucentipollis Khlonova – Srivastava,
p. 546.
Remarks. Following Braman (2013, p. 13), Translucen -
ti pollis is considered to comprise isopolar triprojectate
pollen with reduced polar projections and a more or less
smooth surface.
Translucentipollis contiguus (Tschudy 1969)
Braman 2013
(Plate 17, fig. 13)
Remarks. The specimen illustrated has very fine granu-
lae to short and delicate rugulae that are barely
discernible in optical section. We consider that this wall
surface fits within the granular to scabrate range of
Translucentipollis contiguous.
Age. LO: not determined. Not plotted.
Genus Wodehouseia Stanley 1961
Type. Stanley 1961, plate 1, figs 1–3, as Wodehouseia
spinata.
1961 Wodehouseia Stanley, p. 157.
Remarks. Wodehouseia is included in the ‘oculata’ pol -
len, which Khlonova (1962, p. 306) defined as pollen
grains with two pairs of apertures that are located close
to the tips of the two long sides. Wiggins (1976) gave a
thorough review of various genera in the group, includ-
ing Wodehouseia.
Wodehouseia spinata Stanley 1961
(Plate 20, fig. 3)
1961 Wodehouseia spinata Stanley, p. 157, 158,
160, plate 1, figs 1–12.
Age. LO: latest Maastrichtian.
Genus Zlivisporis Pacltová 1961
Type. Pacltová 1961, plate 2, figs 1–3, as Zlivisporis
blanensis.
1961 Zlivisporis Pacltová, p. 40.
1961 Seductisporites Khlonova, p. 56.
1962 Rouseisporites Pocock, p. 52.
Remarks. The reticulum developed on the distal surface
is never fine and is often incomplete.
Zlivisporis spp.
(Plate 20, figs 4–8)
Age. LO: early Rupelian.
Remarks. Specimens of Zlivisporis from the Labrador–
Baffin Seaway region are variable, especially in the
nature of the surface reticulum, and are thus difficult to
speciate.
Genus Zonalapollenites Pflug in Thomson & Pflug
1953
Type. Potonié 1931, fig. 2, as Sporonites igniculus.
1934 Tsugaepollenites Potonié & Venitz, p. 17;
name not validly published.
1958 Zonalapollenites Pflug in Thomson & Pflug,
p. 66.
1958 Tsugaepollenites Potonié & Venitz ex Potonié,
p. 48.
Remarks. This genus is characterised by the numerous
small, distinct equatorial sacs or vesiculae. Sometimes
the polar regions may be covered with greatly reduced
sacs. Jansonius & Hills (1976, card 3265) recognised
that Tsugaepollenites is an obligate junior synonym of
Zonalapollenites.
Zonalapollenites igniculus (Potonié 1931)
Thomson & Pflug 1953
(Plate 20, figs 9–11)
1931 Sporonites igniculus Potonié, p. 556, fig. 2.
1934 Tsugaepollenites igniculus (Potonié) – Potonié
& Venitz, p. 17; name not validly published.
1953 Zonalapollenites igniculus (Potonié ex Pflug) –
Thomson & Pflug, p. 66.
Age. LO: latest Gelasian.
Bulletin36.qxp_Bulletin 36 19/12/16 13.40 Side 87
88
Fungal elements
Remarks. All fungal spores are grouped together but
several different morphologies are recorded, including
those shown in the accompanying illustrations (Plate
20, figs 12–20). Fungal spores occur sporadically
throughout the sections studied, but peaks were noted
in the Lutetian and Ypresian.
Acknowledgements
During the gestation of this paper of over a decade, we
have had the unwavering support of our two institu-
tions, the Geological Survey of Denmark and Green -
land (GEUS) and the Geological Survey of Canada
(Atlantic) (GSCA), part of the Earth Sciences Sector
(ESS) of Natural Resources Canada. We are also grate-
ful for fruitful discussions with, and feedback from,
Kate Dickie of GSCA, Lotte M. Larsen and Gunver K.
Pedersen of GEUS, Stefan Piasecki of the Geological
Museum, Natural History Museum of Denmark,
University of Copenhagen. We are grateful to Jennifer
Galloway, Martin Pearce, Jim Riding and Art Sweet for
helpful reviews that led to important improvements of
the manuscript. We also extend our thanks Bernie
Crilley and Bill MacMillan (GSC), and Annette Ryge
and Dorthe Salomonsen (GEUS) for processing
samples.
Bulletin36.qxp_Bulletin 36 19/12/16 13.40 Side 88
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Scale
The scale bar represents 20 μm in all plates.
Explanation
In the following plates, details concerning the figured
specimens are presented in a standardised, abbreviated
form grouped in four categories:
Location
Locality: mainly offshore wells, but also on -•
shore boreholes and outcrop sections.
Sample depth (in metres) in wells and boreholes;•
sample number in outcrop sections, unless spe -
cified otherwise.
Sample
Type: cs: cuttings sample. os: outcrop section•
sample. sw: sidewall core sample. cc: conven-
tional core sample.
Sample number or processing number: All ‘P•
numbers’ (e.g. P39553) nos are sample numbers
from Cana dian wells processed in Canada; the
remainder are GGU/GEUS processing numbers
of both Canadian and Greenlandic material.
Slide number/letter.•
Optical parameters
Microscope identification (relating to follow-•
ing Vernier scale coordinates): A: Leitz Dialux
22 microscope 512 742/057691 at the Geol -
ogical Survey of Denmark and Greenland
(GEUS). B: Leica DM 2000 331596-092011 at
GEUS. C: Zeiss Axioplan 2 microscope, serial
no. 310243 at GSC Atlantic, Dartmouth, Nova
Scotia. D: Zeiss Photomicroscope, serial no.
67750 at GSC Atlantic, Dartmouth, Nova
Scotia. E: Leitz DM RB (RS232C) at GEUS.
Vernier coordinates.•
England Finder coordinates.•
Lens magnification and type: bf: bright field.•
pc: phase contrast.
Repositories
CNLOPB: slide curated in the collection of•
the Canada – Newfoundland and Labrador
Offshore Petroleum Board, St. John’s, New -
foundland, Canada (no numbers).
CNSOPB: slide curated in the collection of•
the Canada – Nova Scotia Offshore Petroleum
Board, Dartmouth, Nova Scotia, Canada (no
numbers).
MGUH: slide curated in the type collection of•
the Geological Museum of the University of
Copenhagen (at the time of writing, this mate-
rial is on long-term loan to GEUS, Øster Vold -
gade 10, DK-1350 Copenhagen K, Denmark).
GSC: slide curated in the National Collection•
of Type Invertebrate and Plant Fossils, Geol -
ogical Survey of Canada, 601 Booth Street, Ot ta -
wa, Ontario, Canada K1A 0E8 (at the time of
writing, this material is on long-term loan to
GSC Atlantic, Bedford Institute of Oceano -
graphy, Dartmouth, Nova Scotia, Canada B2Y
4A2).
Statoil: slides are stored at the offices of the•
Statoil petroleum company in Stavanger, Nor -
way (no numbers).
All nomenclatural types are curated at either•
the Geological Museum of the University of
Co pen hagen or at the Geological Survey of
Canada, as indicated above.
Previous publication
The following figures were previously published (in
black-and-white) in Nøhr-Hansen (2003): Plate 2, fig.
12; Plate 11, figs 18,19; Plate 15, figs 9–12; Plate 17,
fig. 2. In addition, Plate 6, fig. 1 was previously published
in colour in Nøhr-Hansen (1996).
Plates
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104
Fig. 1. Achilleodinium biformoides, optical section.
Location: North Leif I-05, 2040‒2050 m. Sample: cs, YD17597, 3.
Optical parameters: E, 34.6 × 17.7, K33/1, × 60 bf.
Repository: GSC 138151.
Fig. 2. Achilleodinium biformoides, same specimen as fig. 1,
focussed on ventral surface.
Location: North Leif I-05, 2040‒2050 m. Sample: cs, YD17597, 3.
Optical parameters: E, 34.6 × 17.7, K33/1, × 60 bf.
Repository: GSC 138152.
Fig. 3. Achilleodinium biformoides, dorsal surface.
Location: South Labrador N-79, 1920‒1930 m. Sample: cs, P39782, 1.
Optical parameters: C, 192 × 1020, T31/4, × 50 bf.
Repository: GSC 138115.
Fig. 4. Adnatosphaeridium vittatum, dorsal surface.
Location: Karlsefni A-13, 1627.65–1636.80 m. Sample: cs, P39577, 1.
Optical parameters: D, 179 × 866, R29/3, × 40 pc.
Repository: GSC 138013.
Fig. 5. Alisocysta margarita, dorsal surface.
Location: North Leif I-05, 2310‒2320 m. Sample: cs, YD17606, 2.
Optical parameters: E, 28.5 × 6.9, U27/3, × 60 bf.
Repository: MGUH 31266.
Fig. 6. Alisocysta circumtabulata, ventral view of ventral surface.
Location: South Labrador N-79, 3060‒3070 m. Sample: cs, P39819, 1.
Optical parameters: C, 166 × 1026, R32/0–2, × 50 bf.
Repository: GSC 138124.
Fig. 7. Alisocysta circumtabulata, same specimen as fig. 6, focussed
on dorsal surface.
Location: South Labrador N-79, 3060‒3070 m. Sample: cs, P39819, 1.
Optical parameters: C, 166 × 1026, R32/0–2, × 50 bf.
Repository: GSC 138124
Fig. 8. Adnatosphaeridium vittatum, apical view.
Location: South Labrador N-79, 2310‒2320 m. Sample: cs, P39795, 1.
Optical parameters: C, 184 × 1028, S32/4, × 50 bf.
Repository: GSC 138120.
Fig. 9. Alisocysta margarita, dorsal view.
Location: North Leif I-05, 2370‒2380 m. Sample: cs, YD17608, 4.
Optical parameters: E, 38.9 × 13.6, O37/2, × 60 bf.
Repository: GSC 138161.
Fig. 10. Alterbidinium acutulum, dorsal view of dorsal surface.
Location: Karlsefni A-13, 1106.44–1115.58 m. Sample: cs, P39558, 1.
Optical parameters: D, 192 × 918, T34/2, × 40 pc.
Repository: GSC 138010.
Fig. 11. Alterbidinium acutulum, dorsal surface.
Location: Skolp E-07, 1295 m. Sample: cs, YD15594, 3.
Optical parameters: A, 49.0 × 113.2, C50/1, × 60 bf.
Repository: MGUH 31267.
Fig. 12. Alterbidinium biaperturum, dorsal view of dorsal surface.
Location: Skolp E-07, 1070 m. Sample: cs, YD15580, 3.
Optical parameters: A, 19.6 × 96.4, U19/2, × 60 bf.
Repository: MGUH 31268.
Fig. 13. Alterbidinium biaperturum, same specimen as fig. 12,
dorsal view of ventral surface.
Location: Skolp E-07, 1070 m. Sample: cs, YD15580, 3.
Optical parameters: A, 19.6 × 96.4, U19/2, × 60 bf.
Repository: MGUH 31268.
Fig. 14. Alterbidinium biaperturum, dorsal view of ventral surface.
Location: Skolp E-07, 1250 m. Sample: cs, YD15592, 3.
Optical parameters: A, 25.2 × 98.0, S25/3, × 60 bf.
Repository: MGUH 31269.
Fig. 15. Alterbidinium? bicellulum, ventral view.
Location: Gjoa O-37, 1620 m. Sample: cs, YD16074, 2.
Optical parameters: A, 16.2 × 100.0, R16/1, × 60 bf.
Repository: MGUH 31270.
Fig. 16. Alterbidinium? bicellulum, ventral view.
Location: Gjoa O-37, 1620 m. Sample: cs, YD16074, 2.
Optical parameters: A, 29.2 × 93.0, Y29/0, × 60 bf.
Repository: MGUH 31271.
Fig. 17. Alterbidinium ioannidesii, dorsal view of ventral surface.
Location: Skolp E-07, 2390 m. Sample: cs, YD15666, 2.
Optical parameters: A, 20.1 × 113.2, B20/3, × 60 bf.
Repository: MGUH 31272.
Fig. 18. Alterbidinium varium, ventral view.
Location: Skolp E-07, 1715 m. Sample: cs, JEH15622, 3.
Optical parameters: A, 40.0 × 103.7, M40/4, × 60 bf.
Repository: MGUH 31273.
Fig. 19. Alterbidinium varium, dorsal view.
Location: Skolp E-07, 1715 m. Sample: cs, JEH15622, 5.
Optical parameters: A, 26.0 × 101.6, P26/3, × 60 bf.
Repository: CNLOPB.
Fig. 20. Apectodinium homomorphum, dorsal view.
Location: North Leif I-05, 2010‒2020 m. Sample: cs, YD17596, 2.
Optical parameters: E, 34.1 × 12.8, O33/3, × 60 bf.
Repository: MGUH 31274.
Plate 1
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105
1
9
5
13
17
2
10
6
14
18
3
11
7
15
19
4
12
8
16
20
Plate 1
Plate 1
Bulletin36.qxp_Bulletin 36 19/12/16 13.40 Side 105
106
Fig. 1. Apectodinium parvum, ventral view.
Location: North Leif I-05, 2100‒2110 m. Sample: cs, YD17599, 3.
Optical parameters: E, 22.6× 16.8, K20/4, × 60 bf.
Repository: GSC 138154.
Fig. 2. Apectodinium parvum, dorsal view.
Location: North Leif I-05, 2070‒2080 m. Sample: cs, YD17598, 4.
Optical parameters: E, 20.3 × 23.3, D18/0, × 60 bf.
Repository: MGUH 31275.
Fig. 3. Apectodinium parvum, dorsal view of dorsal surface.
Location: Rut H-11, 3335‒3345 m. Sample: cs, P39405, 1.
Optical parameters: D, 121 × 1034, M46/2, × 40 pc.
Repository: GSC 137967.
Fig. 4. Apectodinium quinquelatum, dorsal view of ventral surface.
Location: North Leif I-05, 2250‒2260 m. Sample: cs, YD17604, 3.
Optical parameters: E, 55.2 × 18.9, G54/4, × 60 bf.
Repository: MGUH 31276.
Fig. 5. Aptea polymorpha, dorsal view.
Location: Roberval K-92, 3160‒3170 m. Sample: cs, P2008177, 1.
Optical parameters: C, 79 × 980, H27/7, × 50 bf.
Repository: GSC 137910.
Fig. 6. Apteodinium australiense, dorso-ventral view.
Location: Karlsefni A-13, 969.28–978.42 m. Sample: cs, P39553, 1.
Optical parameters: D, 167 × 898, Q32/4, × 40 pc.
Repository: GSC 138005.
Fig. 7. Apteodinium spiridoides, right lateral view.
Location: Karlsefni A-13, 2505.49–2514.63 m. Sample: cs, P39608, 1.
Optical parameters: D, 171 × 990, R42/0, × 40 pc.
Repository: GSC 138037.
Fig. 8. Apteodinium spiridoides, right lateral view.
Location: Rut H-11, 725‒735 m. Sample: cs, P39318, 1.
Optical parameters: D, 78 × 913, G34/0‒3, × 40 pc.
Repository: GSC 137919.
Fig. 9. Areoligera circumsenonensis, ventral view of dorsal surface.
Location: Karlsefni A-13, 2615.22–2624.36 m. Sample: cs, P39612, 1.
Optical parameters: D, 183 × 990, S42/1‒3, × 40 pc.
Repository: GSC 138040.
Fig. 10. Areoligera gippingensis, dorso-ventral view.
Location: Karlsefni A-13, 3822.24–3831.38 m. Sample: cs, P39655, 1.
Optical parameters: D, 143 × 930, O36/1, × 40 pc.
Repository: GSC 138054.
Fig. 11. Areoligera gippingensis, dorso-ventral view.
Location: Gjoa O-37, 2240 m. Sample: cs, YD16095, 2.
Optical parameters: E, 52.3 × 14.2, M51/4, × 60 bf.
Repository: MGUH 31277.
Fig. 12. Areosphaeridium diktyoplokum, dorso-ventral view.
Location: Hellefisk-1, 723 m. Sample: sw, YD13709, 2.
Optical parameters: A, 38.9 × 108.4, H39/4, × 60 bf.
Repository: MGUH 26473.
Fig. 13. Atopodinium cf. haromense, dorso-ventral view.
Location: Skolp E-07, 1550 m. Sample: cs, JEH15611, 5.
Optical parameters: A, 35.0 × 95.5, V35/1, × 60 bf.
Repository: MGUH 31278.
Fig. 14. Axiodinium augustum, dorso-ventral view.
Location: North Leif I-05, 2250‒2260 m. Sample: cs, YD17604, 3.
Optical parameters: E, 43.0 × 21.6, E42/1, × 60 bf.
Repository: MGUH 31279.
Fig. 15. Axiodinium augustum, dorso-ventral view, same specimen
as fig. 14, focussed on antapical part.
Location: North Leif I-05, 2250‒2260 m. Sample: cs, YD17604, 3.
Optical parameters: E, 43.0 × 21.6, E42/1, × 60 bf.
Repository: MGUH 31279.
Fig. 16. Batiacasphaera micropapillata, dorso-ventral view.
Location: Rut H-11, 845‒855 m. Sample: cs, P39322, 1.
Optical parameters: D, 93 × 1020, J45/0, × 40 pc.
Repository: GSC 137924.
Fig. 17. Batioladinium jaegeri, ventral view of ventral surface.
Location: Skolp E-07, 1655 m. Sample: cs, JEH15618, 3.
Optical parameters: A, 45.7 × 107.9, H46/4, × 60 bf.
Repository: MGUH 31281.
Fig. 18. Callaiosphaeridium asymmetricum, apical-antapical view.
Location: Skolp E-07, 1535 m. Sample: cs, JEH15610, 3.
Optical parameters: A, 33.7 × 98.8, R34/3, × 60 bf.
Repository: MGUH 31282.
Fig. 19. Cerebrocysta bartonensis, left lateral view.
Location: Ogmund E-72, 1336 m. Sample: sw, JEH15552, 2.
Optical parameters: A, 39.8 × 107.7, H40/4, × 60 bf.
Repository: MGUH 31283.
Fig. 20. Cerebrocysta bartonensis, left lateral view, same specimen
as fig. 19, focussed on operculum, within cyst.
Location: Ogmund E-72, 1336 m. Sample: sw, JEH15552, 2.
Optical parameters: A, 39.8 × 107.7, H40/4, × 60 bf.
Repository: MGUH 31283.
Plate 2
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107
1
9
13
17
2
10
14
18
3
11
7
15
4
12
8
16
20
Plate 2
65
19
Plate 2
Bulletin36.qxp_Bulletin 36 19/12/16 13.40 Side 107
108
Fig. 1. Cerodinium diebelii, dorso-ventral view.
Location: Roberval K-92, 3160‒3170 m. Sample: cs, P2008177, 1.
Optical parameters: C, 130 × 918, N20/0, × 40 bf.
Repository: GSC 137909.
Fig. 2. Cerodinium diebelii, ventral view of dorsal surface.
Location: Roberval K-92, 2500‒2510 m. Sample: cs, P19229, 1.
Optical parameters: C, 100 × 1018, K31/0–2, × 50 bf.
Repository: GSC 137905.
Fig. 3. Cerodinium diebelii, dorsal view of dorsal surface.
Location: North Leif I-05, 2370‒2380 m. Sample: cs, YD17608, 2.
Optical parameters: E, 39.6 × 16.1, L38/2, × 60 bf.
Repository: MGUH 31284.
Fig. 4. Cerodinium diebelii, dorso-ventral view.
Location: North Leif I-05, 2520‒2530 m. Sample: cs, YD17613, 3.
Optical parameters: E, 23.8 × 18.7, H22/3, × 60 bf.
Repository: MGUH 31285.
Fig. 5. Cannosphaeropsis passio, oblique antapical view.
Location: Qulleq-1, 1847 m. Sample: sw, YD14581, 4.
Optical parameters: A, 38.4 × 108.0, J38/2, × 40 bf.
Repository: Statoil.
Fig. 6. Cerebrocysta magna, oblique right lateral view.
Location: Qulleq-1, 1862.1 m. Sample: sw, YD14587, 3.
Optical parameters: A, 29.8 × 100.7, Q30/3, × 60 bf.
Repository: Statoil/MGUH 26505.
Fig. 7. Cerodinium glabrum, dorsal surface.
Location: Gilbert F-53, 3420‒3430 m. Sample: cs, P17040, 1.
Optical parameters: C, 158 × 1010, L30/2, × 50 bf.
Repository: GSC 137887.
Fig. 8. Cerodinium glabrum, dorsal surface.
Location: Gjoa O-37, 2400 m. Sample: cs, YD16097, 2.
Optical parameters: E, 21.4 × 25.3, B19/2, × 60 bf.
Repository: MGUH 31286.
Fig. 9. Cerodinium kangiliense, dorsal view of dorsal surface.
Location: Skolp E-07, 1460 m. Sample: cs, YD15605, 3.
Optical parameters: A, 52.0 × 102.8, N53/3, × 60 bf.
Repository: MGUH 31287.
Fig. 10. Cerodinium kangiliense, dorsal view, same specimen as fig.
9, focussed on ventral surface.
Location: Skolp E-07, 1460 m. Sample: cs, YD15605, 3.
Optical parameters: A, 52.0 × 102.8, N53/3, × 60 bf.
Repository: MGUH 31287.
Fig. 11. Cerodinium striatum, dorso-ventral view of dorsal surface.
Location: Gilbert F-53, 2410‒2420 m. Sample: cs, P39493, 1.
Optical parameters: C, 70 × 1040, G34/1, × 50 bf.
Repository: GSC 137983.
Fig. 12. Cerodinium striatum, dorso-ventral view.
Location: Rut H-11, 2765‒2775 m. Sample: cs, P39386, 1.
Optical parameters: D, 65 × 869, F29/2, × 40 pc.
Repository: GSC 137955.
Fig. 13. Cerodinium speciosum, dorso-ventral view of dorsal surface.
Location: Hekja O-71, 4510‒4520 m. Sample: cs, P20645, 1.
Optical parameters: C, 50 × 1007, E30/0, × 50 bf.
Repository: GSC 137915.
Fig. 14. Chatangiella madura, dorsal view of dorsal surface.
Location: Skolp E-07, 1580 m. Sample: cs, JEH15613, 3.
Optical parameters: A, 31.0 × 112.8, C31/4, × 60 bf.
Repository: MGUH 31288.
Fig. 15. Chatangiella madura, dorsal view, same specimen as fig.
14, focussed on optical section.
Location: Skolp E-07, 1580 m. Sample: cs, JEH15613, 3.
Optical parameters: A, 31.0 × 112.8, C31/4, × 60 bf.
Repository: MGUH 31288.
Fig. 16. Chatangiella tripartita, dorsal view of dorsal surface.
Location: South Labrador N-79, 1230‒1240 m. Sample: cs, P39761, 1.
Optical parameters: C, 178 × 988, S28/0–2, × 50 bf.
Repository: GSC 138081.
Fig. 17. Chiropteridium galea, dorsal view.
Location: Karlsefni A-13, 1051.57–1060.72 m. Sample: cs, P39556, 1.
Optical parameters: D, 228 × 1029, X46/1, × 40 pc.
Repository: GSC 138008.
Fig. 18. Chiropteridium galea, ventral view of ventral surface.
Location: Karlsefni A-13, 1079.01–1088.15 m. Sample: cs, P39557, 1.
Optical parameters: D, 170 × 991, R42/1, × 40 pc.
Repository: GSC 138009.
Fig. 19. Chiropteridium galea, ventral view.
Location: Karlsefni A-13, 1161.30–1170.45 m. Sample: cs, P39560, 1.
Optical parameters: D, 172 × 889, R31/2, × 40 pc.
Repository: GSC 138011.
Fig. 20. Chatangiella tripartita, ventral view of dorsal surface.
Location: Karlsefni A-13, 2267.74–2276.88 m. Sample: cs, P39599, 1.
Optical parameters: D, 192 × 926, T35/0, × 40 pc.
Repository: GSC 138029.
Plate 3
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109
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Plate 3
Plate 3
Bulletin36.qxp_Bulletin 36 19/12/16 13.40 Side 109
110
Fig. 1. Chiropteridium gilbertii sp. nov., dorso-ventral view.
Location: South Labrador N-79, 1440‒1450 m. Sample: cs, P39768, 1.
Optical parameters: C, 56 × 1028, E32/4, × 50 bf.
Repository: GSC 138112.
Fig. 2. Chiropteridium gilbertii sp. nov., ventral view of dorsal surface.
Location: South Labrador N-79, 1560‒1570 m. Sample: cs, P39772, 1.
Optical parameters: C, 168 × 1031, R33/1, × 50 bf.
Repository: GSC 138113.
Fig. 3. Chiropteridium gilbertii sp. nov., ventral view, same speci-
men as fig. 2, focussed on ventral surface.
Location: South Labrador N-79, 1560‒1570 m. Sample: cs, P39772, 1.
Optical parameters: C, 168 × 1031, R33/1, × 50 bf.
Repository: GSC 138113.
Fig. 4. Chiropteridium gilbertii sp. nov., holotype, ventral view.
Location: Gilbert F-53, 1600‒1610 m. Sample: cs, P39466, 1.
Optical parameters: C, 174 × 1060, R36/3, × 50 bf.
Repository: GSC 137976.
Fig. 5. Chlamydophorella nyei, left lateral view.
Location: South coast Bylot Island, section B, 1.8 m above base (sample
HFB09-17A). Sample: os, P5148-4, C.
Optical parameters: C, 193 × 901, H57/0–1, × 50 bf.
Repository: GSC 138130.
Fig. 6. Chlamydophorella cf. nyei, dorso-ventral view.
Location: Ogmund E-72, 1620 m. Sample: cs, YD15740, 5.
Optical parameters: A, 19.5 × 110.5, F19/2, × 60 bf.
Repository: GSC 138145.
Fig. 7. Chlamydophorella cf. nyei, right lateral view.
Location: North Leif I-05, 2670–2680 m. Sample: cs, YD17618, 2.
Optical parameters: E, 18.6 × 19.5, G16/2, × 60 bf.
Repository: MGUH 31289.
Fig. 8. Chlamydophorella cf. nyei, dorso-ventral view.
Location: Skolp E-07, 1685 m. Sample: cs, JEH15620, 3.
Optical parameters: B, 30.47 × 5.36, X32/1, × 60 bf.
Repository: MGUH 31290.
Fig. 9. Charlesdowniea coleothrypta, ventral view of ventral surface.
Location: Gilbert F-53, 2050‒2060 m. Sample: cs, P39481, 1.
Optical parameters: C, 57 × 1014, E31/3, × 50 bf.
Repository: GSC 137981.
Fig. 10. Cleistosphaeridium elegantulum sp. nov., apical view.
Location: Karlsefni A-13, 2203.73–2212.87 m. Sample: cs, P39597, 1.
Optical parameters: D, 122 × 866, M29/0‒1, × 40 pc.
Repository: GSC 138026.
Fig. 11. Cleistosphaeridium elegantulum sp. nov., oblique apical view.
Location: Karlsefni A-13, 2203.73–2212.87 m. Sample: cs, P39597, 1.
Optical parameters: D, 230 × 894, X32/0, × 40 pc.
Repository: GSC 138027.
Fig. 12. Cleistosphaeridium elegantulum sp. nov., holotype, dorso-
ventral view.
Location: Karlsefni A-13, 2286.03–2295.17 m. Sample: cs, P39600, 1.
Optical parameters: D, 179 × 1017, S44/2, × 40 pc.
Repository: GSC 138033.
Fig. 13. Cleistosphaeridium palmatum sp. nov., holotype, dorsal
view of optical section.
Location: Roberval K-92, 2450‒2460 m. Sample: cs, P17706, 1.
Optical parameters: C, 69 × 1006, G30/0–2, × 50 bf.
Repository: GSC 137897.
Fig. 14. Cleistosphaeridium palmatum sp. nov., holotype, dorsal
view, focussed on dorsal surface.
Location: Roberval K-92, 2450‒2460 m. Sample: cs, P17706, 1.
Optical parameters: C, 69 × 1006, G30/0–2, × 50 bf.
Repository: GSC 137897.
Fig. 15. Cleistosphaeridium palmatum sp. nov., dorso-ventral view.
Location: Rut H-11, 3005‒3015 m. Sample: cs, P39394, 1.
Optical parameters: D, 141 × 905, O33/1, × 40 pc.
Repository: GSC 137960.
Fig. 16. Cleistosphaeridium palmatum sp. nov., dorso-ventral
view.
Location: Rut H-11, 3275‒3285 m. Sample: cs, P39403, 1.
Optical parameters: D, 140 × 925, N35/3‒4, × 40 pc.
Repository: GSC 137966.
Fig. 17. Chytroeisphaeridia hadra sp. nov., dorsal surface showing
the archaeopyle.
Location: Roberval K-92, 1700‒1710 m. Sample: cs, P17681, 1.
Optical parameters: C, 48 × 1017, E30/2, × 50 bf.
Repository: GSC 137888.
Fig. 18. Chytroeisphaeridia hadra, sp. nov., holotype, optical
section view.
Location: Roberval K-92, 3120‒3140 m. Sample: cs, P17728, 1.
Optical parameters: C, 195 × 1098, U40/1, × 50 bf.
Repository: GSC 137902.
Fig. 19. Cleistosphaeridium diversispinosum, dorso-ventral view.
Location: Karlsefni A-13, 969.28–978.42 m. Sample: cs, P39553, 1.
Optical parameters: D, 163 × 971, Q40/0, × 40 pc.
Repository: GSC 138004.
Fig. 20. Cleistosphaeridium diversispinosum, operculum.
Location: Karlsefni A-13, 2203.73–2212.87 m. Sample: cs, P39597, 1.
Optical parameters: D, 115 × 972, L40/0, × 40 pc.
Repository: GSC 138025.
Plate 4
Bulletin36.qxp_Bulletin 36 19/12/16 13.40 Side 110
111
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Plate 4
Plate 4
Bulletin36.qxp_Bulletin 36 19/12/16 13.40 Side 111
112
Fig. 1. Cleistosphaeridium polypetellum, dorso-ventral view.
Location: Karlsefni A-13, 2944.40–2953.55 m. Sample: cs, P39624, 1.
Optical parameters: D, 215 × 1017, V44/4, × 40 pc.
Repository: GSC 138047.
Fig. 2. Cleistosphaeridium polypetellum.
Location: Karlsefni A-13, 3054.13–3063.28 m. Sample: cs, P39628, 1.
Optical parameters: D, 151 × 896, P32/1‒2, × 40 pc.
Repository: GSC 138048.
Fig. 3. Cordosphaeridium delimurum, dorso-ventral view.
Location: Karlsefni A-13, 2807.24–2816.39 m. Sample: cs, P39619, 1.
Optical parameters: D, 166 × 976, Q40/0‒4, × 40 pc.
Repository: GSC 138046.
Fig. 4. Cordosphaeridium delimurum, dorso-ventral view, same
specimen as fig. 3.
Location: Karlsefni A-13, 2807.24–2816.39 m. Sample: cs, P39619, 1.
Optical parameters: D, 166 × 976, Q40/0‒4, × 40 pc.
Repository: GSC 138046.
Fig. 5. Cordosphaeridium fibrospinosum, oblique apical view.
Location: Karlsefni A-13, 3989.88–3999.02 m. Sample: cs, P39661, 1.
Optical parameters: D, 122 × 980, M41/0‒1, × 40 pc.
Repository: GSC 138060.
Fig. 6. Cordosphaeridium fibrospinosum, dorsal view.
Location: Karlsefni A-13, 3989.88–3999.02 m. Sample: cs, P39661, 1.
Optical parameters: D, 126 × 897, M32/0, × 40 pc.
Repository: GSC 138061.
Fig. 7. Cordosphaeridium cantharellus, right lateral view.
Location: Rut H-11, 755‒765 m. Sample: cs, P39319, 1.
Optical parameters: D, 110 × 978, L40/2, × 40 pc.
Repository: GSC 137922.
Fig. 8. Cordosphaeridium funiculatum, left lateral view.
Location: Ralegh N-18, 1525 m. Sample: cs, YD16219, 2.
Optical parameters: E, 43.3 × 23.8, C42/1, × 60 bf.
Repository: MGUH 31291.
Fig. 9. Cordosphaeridium gracile, dorsal surface.
Location: Karlsefni A-13, 3081.57–3090.71 m. Sample: cs, P39629, 1.
Optical parameters: D, 171 × 1030, R46/0, × 40 pc.
Repository: GSC 138049.
Fig. 10. Cordosphaeridium inodes, dorsal surface.
Location: Karlsefni A-13, 3931.97–3941.11 m. Sample: cs, P39659, 1.
Optical parameters: D, 97 × 910, J34/1‒3, × 40 pc.
Repository: GSC 138059.
Fig. 11. Dapsilidinium pseudoinsertum sp. nov., dorso-ventral view.
Location: Karlsefni A-13, 2697.51–2706.66 m. Sample: cs, P39615, 1.
Optical parameters: D, 98 × 946, J37/4, × 40 pc.
Repository: GSC 138043.
Fig. 12. Dapsilidinium pseudoinsertum sp. nov., holotype, dorso-
ventral view.
Location: Rut H-11, 2825‒2835 m. Sample: cs, P39388, 1.
Optical parameters: D, 196 × 1051, U48/1‒2, × 40 pc.
Repository: GSC 137957.
Fig. 13. Dapsilidinium pseudocolligerum.
Location: Gilbert F-53, 2020‒2030 m. Sample: cs, P39480, 1.
Optical parameters: C, 45 × 1008, D30/0–4, × 50 bf.
Repository: GSC 137980.
Fig. 14. Dapsilidinium simplex, dorso-ventral view.
Location: Rut H-11, 3845‒3855 m. Sample: cs, P39421, 1.
Optical parameters: D, 165 × 921, Q35/3, × 40 pc.
Repository: GSC 137974.
Fig. 15. Deflandrea denticulata, dorso-ventral view of dorsal surface.
Location: Karlsefni A-13, 4038.65–4047.79 m. Sample: cs, P39663, 1.
Optical parameters: D, 165 × 1014, Q44/0, × 40 pc.
Repository: GSC 138065.
Fig. 16. Deflandrea denticulata, dorso-ventral view of ventral surface.
Location: Karlsefni A-13, 4038.65–4047.79 m. Sample: cs, P39663, 1.
Optical parameters: D, 60 × 1068, E50/3, × 40 pc.
Repository: GSC 138064.
Fig. 17. Deflandrea borealis sp. nov., dorso-ventral view.
Location: Karlsefni A-13, 2697.51–2706.66 m. Sample: cs, P39615, 1.
Optical parameters: D, 169 × 929, Q35/4, × 40 pc.
Repository: GSC 138044.
Fig. 18. Deflandrea borealis sp. nov., dorso-ventral view.
Location: Rut H-11, 2915‒2925 m. Sample: cs, P39391, 1.
Optical parameters: D, 198 × 883, U31/1, × 40 pc.
Repository: GSC 137959.
Fig. 19. Deflandrea borealis sp. nov., holotype, dorso-ventral view.
Location: Rut H-11, 3095‒3105 m. Sample: cs, P39397, 1.
Optical parameters: D, 189 × 986, T41/2, × 40 pc.
Repository: GSC 137964.
Fig. 20. Deflandrea borealis sp. nov., dorso-ventral view.
Location: Gilbert F-53, 1810‒1820 m. Sample: cs, P39473, 1.
Optical parameters: C, 175 × 1014, R31/3, × 50 bf.
Repository: GSC 137977.
Plate 5
Bulletin36.qxp_Bulletin 36 19/12/16 13.40 Side 112
113
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Plate 5
Plate 5
Bulletin36.qxp_Bulletin 36 19/12/16 13.40 Side 113
114
Fig. 1. Deflandrea galeata, dorso-ventral view of dorsal surface.
Location: Annertuneq, 388 m, GGU366591. Sample: os, C402-G, 5.
Optical parameters: A, 56.1 × 112.8, B556/4, × 60 bf.
Repository: MGUH 23924.
Fig. 2. Deflandrea majae, ventral view of ventral surface.
Location: Skolp E-07, 1055 m. Sample: cs, YD15579, 4.
Optical parameters: A, 45.7 × 96.9, T46/4, × 60 bf.
Repository: GSC 138142.
Fig. 3. Deflandrea oebisfeldensis, dorso-ventral view of ventral surface.
Location: Hekja O-71, 3360 m. Sample: cs, JEH16039, 3.
Optical parameters: E, 32.1 × 5.6, W31/1, × 60 bf.
Repository: MGUH 31292.
Fig. 4. Deflandrea phosphoritica, dorso-ventral view.
Location: Snorri J-90, 2249.45‒2256.6 m. Sample: cs, P9747, 10.
Optical parameters: C, 220 × 1035, W33/0, × 50 bf.
Repository: GSC 138136.
Fig. 5. Diphyes brevispinum, dorso-ventral view.
Location: South Labrador N-79, 1920‒1930 m. Sample: cs, P39782, 1.
Optical parameters: C, 118 × 1030, M32/2–M33/1, × 50 bf.
Repository: GSC 138114.
Fig. 6. Diphyes brevispinum, dorso-ventral view.
Location: South Labrador N-79, 1980‒1990 m. Sample: cs, P39784, 1.
Optical parameters: C, 144 × 1042, O34/3, × 50 bf.
Repository: GSC 138117.
Fig. 7. Diphyes ficusoides, dorso-ventral view.
Location: Bjarni O-82, 1905‒1915 m. Sample: cs, P39718, 1.
Optical parameters: C, 130 × 1061, N36/1–2, × 50 bf.
Repository: GSC 138071.
Fig. 8. Diphyes ficusoides, dorso-ventral view.
Location: Rut H-11, 3125‒3135 m. Sample: cs, P39398, 1.
Optical parameters: D, 164 × 971, Q40/0, × 40 pc.
Repository: GSC 137965.
Fig. 9. Disphaerogena carposphaeropsis.
Location: Annertuneq, 451 m, GGU405093. Sample: os, YD11738, 3.
Optical parameters: A, 27.6 × 96.7, U27/2, × 40 bf.
Repository: MGUH 31293.
Fig. 10. Cyclonephelium distinctum, dorsal view of dorsal surface.
Location: Gilbert F-53, 3220‒3230 m. Sample: cs, P39520, 1.
Optical parameters: C, 28 × 1026, B32/3–4, × 50 bf.
Repository: GSC 137990.
Fig. 11. Dinogymnium longicorne.
Location: Skolp E-07, 1895 m. Sample: cs, JEH15633, 4.
Optical parameters: A, 42.9 × 110.5, E43/4, × 60 bf.
Repository: MGUH 31294.
Fig. 12. Diphyes colligerum, dorso-ventral view.
Location: Gilbert F-53, 1990‒2000 m. Sample: cs, P39479, 1.
Optical parameters: C, 194 × 1058, T35/4, × 50 bf.
Repository: GSC 137979.
Fig. 13. Eatonicysta furensis, dorsal view of dorsal surface.
Location: North Leif I-05, 2070‒2080 m. Sample: cs, YD17598, 3.
Optical parameters: E, 19.3 × 15.7, M17/0, × 60 bf.
Repository: GSC 138153.
Fig. 14. Eatonicysta ursulae, dorso-ventral view.
Location: South Labrador N-79, 2220‒2230 m. Sample: cs, P39792, 1.
Optical parameters: C, 56 × 1042, E34/3, × 50 bf.
Repository: GSC 138119.
Fig. 15. Evittosphaerula? foraminosa sp. nov., apical view.
Location: North Leif I-05, 2130‒2140 m. Sample: cs, YD17600, 3.
Optical parameters: E, 23.7 × 10.0, S22/1, × 60 bf.
Repository: GSC 138155.
Fig. 16. Evittosphaerula? foraminosa sp. nov., apical view of apical
surface.
Location: North Leif I-05, 2190‒2200 m. Sample: cs, YD17602, 3.
Optical parameters: E, 36.5 × 26.0, A35/0, × 60 bf.
Repository: GSC 138160.
Fig. 17. Evittosphaerula? foraminosa sp. nov., oblique apical view
of epicyst.
Location: North Leif I-05, 2130‒2140 m. Sample: cs, YD17600, 3.
Optical parameters: E, 32.8 × 20.8, F31/3, × 60 bf.
Repository: GSC 138157.
Fig. 18. Evittosphaerula? foraminosa sp. nov., oblique antapical view.
Location: North Leif I-05, 2130‒2140 m. Sample: cs, YD17600, 3.
Optical parameters: E, 34.2 × 5.7, W32/2, × 60 bf.
Repository: GSC 138158.
Fig. 19. Evittosphaerula? foraminosa sp. nov., holotype, right late -
ral view focussed on right lateral surface.
Location: North Leif I-05, 2130‒2140 m. Sample: cs, YD17600, 3.
Optical parameters: E, 44.9 × 15.5, M43/0, × 60 bf.
Repository: GSC 138159.
Fig. 20. Evittosphaerula? foraminosa sp. nov., holotype, right late -
ral view, focussed on left lateral surface.
Location: North Leif I-05, 2130‒2140 m. Sample: cs, YD17600, 3.
Optical parameters: E, 44.9 × 15.5, M43/0, × 60 bf.
Repository: GSC 138159.
Plate 6
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Plate 6
Plate 6
Bulletin36.qxp_Bulletin 36 19/12/16 13.40 Side 115
116
Fig. 1. Enneadocysta magna, ventral view.
Location: Rut H-11, 1205‒1215 m. Sample: cs, P39334, 1.
Optical parameters: D, 100 × 990, J42/3, × 40 pc.
Repository: GSC 137929.
Fig. 2. Enneadocysta magna, ventral view.
Location: Rut H-11, 1235‒1245 m. Sample: cs, P39335, 1.
Optical parameters: D, 216 × 859, V28/3‒4, × 40 pc.
Repository: GSC 137933.
Fig. 3. Eocladopyxis peniculata.
Location: South Labrador N-79, 2310‒2320 m. Sample: cs, P20171, 1.
Optical parameters: C, 190 × 1100, T41/0–3, × 50 bf.
Repository: GSC 137911.
Fig. 4. Fibrocysta bipolaris, right lateral view.
Location: Roberval K-92, 2390‒2400 m. Sample: cs, P17704, 1.
Optical parameters: C, 33 × 1090, C39/0, × 50 bf.
Repository: GSC 137895.
Fig. 5. Ginginodinium? flexidentatum sp. nov., dorso-ventral view
of ventral surface.
Location: North Leif I-05, 2010‒2020 m. Sample: cs, YD17596, 3.
Optical parameters: E, 24.9 × 21.8, E23/3, × 60 bf.
Repository: GSC 138150.
Fig. 6. Ginginodinium? flexidentatum sp. nov., dorso-ventral view
of dorsal surface.
Location: Ogmund E-72, 1340 m. Sample: cs, YD15727, 3.
Optical parameters: A, 43.4 × 96.5, U44/1, × 60 bf.
Repository: MGUH 31295.
Fig. 7. Ginginodinium? flexidentatum sp. nov., dorso-ventral view
of ventral surface.
Location: Hekja O-71, 3090 m. Sample: cs, JEH16029, 2.
Optical parameters: E, 35.6 × 20.8, F34/0, × 60 bf.
Repository: MGUH 31296.
Fig. 8. Ginginodinium? flexidentatum sp. nov., dorso-ventral view.
Location: Karlsefni A-13, 2807.24–2816.39 m. Sample: cs, P39619, 1.
Optical parameters: D, 146 × 1060, O49/0, × 40 pc.
Repository: GSC 138045.
Fig. 9. Ginginodinium? flexidentatum sp. nov., dorso-ventral view.
Location: North Leif I-05, 2010‒2020 m. Sample: cs, YD17596, 3.
Optical parameters: A, 22.5 × 19.3, H20/0, × 60 bf.
Repository: GSC 138149.
Fig. 10. Ginginodinium? flexidentatum sp. nov., dorso-ventral view.
Location: Rut H-11, 2915‒2925 m. Sample: cs, P39391, 1.
Optical parameters: D, 140 × 1029, O46/1, × 40 pc.
Repository: GSC 137958.
Fig. 11. Ginginodinium? flexidentatum sp. nov., holotype, dorso-
ventral view.
Location: Bjarni O-82, 1815‒1825 m. Sample: cs, P39715, 1.
Optical parameters: C, 36 × 1028, C32/4, × 50 bf.
Repository: GSC 138070.
Fig. 12. Gillinia hymenophora.
Location: Gilbert F-53, 3280‒3290 m. Sample: cs, P39522, 1.
Optical parameters: C, 155 × 1045, P34/3–4, × 50 bf.
Repository: GSC 137991.
Fig. 13. Habibacysta tectata, right lateral view, showing detached
operculum.
Location: Karlsefni A-13, 667.52–676.66 m. Sample: cs, P39542, 1.
Optical parameters: D, 180 × 979, S41/1, × 40 pc.
Repository: GSC 137995.
Fig. 14. Glaphyrocysta exuberans, dorso-ventral view.
Location: North Leif I-05, 1530‒1540 m. Sample: cs, YD15580, 2.
Optical parameters: A, 38.8 × 23.5, C37/4, × 60 bf.
Repository: MGUH 31297.
Fig. 15. Glaphyrocysta divaricata, dorso-ventral view.
Location: Karlsefni A-13, 3438.19–3447.33 m. Sample: cs, P39642, 1.
Optical parameters: D, 194 × 1048, T48/3, × 40 pc.
Repository: GSC 138053.
Fig. 16. Glaphyrocysta divaricata, dorso-ventral view.
Location: Karlsefni A-13, 3438.19–3447.33 m. Sample: cs, P39642, 1.
Optical parameters: D, 188 × 1032, S46/3‒4, × 40 pc.
Repository: GSC 138052.
Fig. 17. Glaphyrocysta retiintexta, ventral view.
Location: Karlsefni A-13, 4038.65–4047.79 m. Sample: cs, P39663, 1.
Optical parameters: D, 220 × 1066, W49/2‒4, × 40 pc.
Repository: GSC 138066.
Fig. 18. Histiocysta palla, ventral view.
Location: Ogmund E-72, 1620 m. Sample: cs, YD15740, 4.
Optical parameters: A, 37.0 × 98.7, R37/4, × 60 bf.
Repository: MGUH 31298.
Fig. 19. Histiocysta palla, oblique right lateral view, same specimen
as fig. 18.
Location: Ogmund E-72, 1620 m. Sample: cs, YD15740, 4.
Optical parameters: A, 37.0 × 98.7, R37/4, × 60 bf.
Repository: MGUH 31298.
Fig. 20. Histiocysta palla, dorsal surface.
Location: Bjarni O-82, 2300‒2310 m. Sample: cs, P39731, 1.
Optical parameters: C, 140 × 996, O29/0, × 50 bf.
Repository: GSC 138073.
Plate 7
Bulletin36.qxp_Bulletin 36 19/12/16 13.40 Side 116
117
1
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Plate 7
Plate 7
Bulletin36.qxp_Bulletin 36 19/12/16 13.40 Side 117
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Fig. 1. Glaphyrocysta texta, oblique lateral view.
Location: North Leif I-05, 2100‒2110 m. Sample: cs, YD17599, 4.
Optical parameters: E, 25.0 × 16.6, L23/2, × 60 bf.
Repository: MGUH 31299.
Fig. 2. Glaphyrocysta vicina, ventral view of ventral surface.
Location: Ralegh N-18, 1725 m. Sample: cs, YD16224, 2.
Optical parameters: E, 45.9 × 7.7, T45/3, × 60 bf.
Repository: MGUH 31300.
Fig. 3. Heterosphaeridium bellii.
Location: Gilbert F-53, 3460‒3470 m. Sample: cs, P39528, 1.
Optical parameters: C, 62 × 1062, F36/0, × 50 bf.
Repository: GSC 137992.
Fig. 4. Heterosphaeridium bellii.
Location: North Leif I-05, 2670‒2680 m. Sample: cs, YD17618, 3.
Optical parameters: E, 43.0 × 7.1, V41/2, × 60 bf.
Repository: GSC 138166.
Fig. 5. Homotryblium tenuispinosum, apical view, focussed on
archaeopyle margin.
Location: Gilbert F-53, 1960‒1970 m. Sample: cs, P39478, 1.
Optical parameters: C, 194 × 983, T28/3, × 50 bf.
Repository: GSC 137978.
Fig. 6. Homotryblium tenuispinosum, apical view, same specimen as
fig. 5, focussed on hypocyst.
Location: Gilbert F-53, 1960‒1970 m. Sample: cs, P39478, 1.
Optical parameters: C, 194 × 983, T28/3, × 50 bf.
Repository: GSC 137978.
Fig. 7. Homotryblium tenuispinosum, dorso-ventral view.
Location: South Labrador N-79, 1920‒1930 m. Sample: cs, P39782, 1.
Optical parameters: C, 40 × 1061, D35/1, × 50 bf.
Repository: GSC 138116.
Fig. 8. Heterosphaeridium difficile, dorso-ventral view.
Location: South Labrador N-79, 3485‒3495 m. Sample: cs, P20210, 1.
Optical parameters: C, 168 × 1128, R43/1‒2, × 50 bf.
Repository: GSC 137912.
Fig. 9. Homotryblium abbreviatum, dorsal view.
Location: Snorri J-90, 2249.45‒2256.6 m. Sample: cs, P9747, 10.
Optical parameters: C, 58 × 1092, F39/1–2, × 50 bf.
Repository: GSC 138138.
Fig. 10. Homotryblium abbreviatum, apical view.
Location: Snorri J-90, 2249.45‒2266.6 m. Sample: cs, P9747, 10.
Optical parameters: C, 50 × 1000, E29/2, × 50 bf.
Repository: GSC 138137.
Fig. 11. Hystrichokolpoma cinctum, dorsal surface.
Location: Karlsefni A-13, 2642.65–2651.79 m. Sample: cs, P39613, 1.
Optical parameters: D, 128 × 920, M35/3, × 40 pc.
Repository: GSC 138041.
Fig. 12. Hapsocysta? benteae.
Location: North Leif I-05, 2310‒2320 m. Sample: cs, YD17606, 4.
Optical parameters: E, 50.8 × 19.9, G50/3, × 60 bf.
Repository: MGUH 31301.
Fig. 13. Hystrichokolpoma globulus, left lateral view of left lateral
surface.
Location: Snorri J-90, 2293,65 m. Sample: sw, P9370, 10.
Optical parameters: C, 181 × 1017, S31/0–4, × 50 bf.
Repository: GSC 138132.
Fig. 14. Hystrichokolpoma globulus, left lateral view, same specimen
as fig. 13, focussed on right lateral surface.
Location: Snorri J-90, 2293,65 m. Sample: sw, P9370, 10.
Optical parameters: C, 181 × 1017, S31/0–4, × 50 bf.
Repository: GSC 138132.
Fig. 15. Hystrichokolpoma globulus, lateral view.
Location: Snorri J-90, 2441.48‒2450.62 m. Sample: cs, P9751, 10.
Optical parameters: C, 102 × 1010, K30/4, × 50 bf.
Repository: GSC 138139.
Fig. 16. Heteraulacacysta porosa, antapical view of antapical surface.
Location: Ralegh N-18, 1525 m. Sample: cs, YD16219, 4.
Optical parameters: E, 36.3 × 20.5, G35/1, × 60 bf.
Repository: CNSOPB.
Fig. 17. Hystrichosphaeridium quadratum sp. nov., holotype, dorsal
surface.
Location: Gilbert F-53, 2770‒2780 m. Sample: cs, P39505, 1.
Optical parameters: C, 113 × 992, L/29/3, × 50 bf.
Repository: GSC 137988.
Fig. 18. Hystrichosphaeridium quadratum sp. nov., holotype,
focussed on ventral surface.
Location: Gilbert F-53, 2770‒2780 m. Sample: cs, P39505, 1.
Optical parameters: C, 113 × 992, L29/3, × 50 bf.
Repository: GSC 137988.
Fig. 19. Hystrichosphaeridium quadratum sp. nov., operculum,
showing pre-apical process.
Location: Karlsefni A-13, 3904.54–3913.68 m. Sample: cs, P39658, 1.
Optical parameters: D, 125 × 999, M43/0‒3, × 40 pc.
Repository: GSC 138057.
Fig. 20. Hystrichosphaeridium tubiferum, lateral view.
Location: Gjoa O-37, 2120 m. Sample: cs, YD16090, 4.
Optical parameters: E, 32.7 × 22.0, E31/0, × 60 bf.
Repository: CNSOPB.
Plate 8
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Plate 8
Plate 8
Bulletin36.qxp_Bulletin 36 19/12/16 13.40 Side 119
120
Fig. 1. Hystrichostrogylon digitus sp. nov., dorso-ventral view.
Location: Karlsefni A-13, 2267.74–2276.88 m. Sample: cs, P39599, 1.
Optical parameters: D, 200 × 878, U30/0, × 40 pc.
Repository: GSC 138030.
Fig. 2. Hystrichostrogylon digitus sp. nov., dorso-ventral view.
Location: Rut H-11, 2375‒2385 m. Sample: cs, P39373, 1.
Optical parameters: D, 167 × 973, Q40/3‒4, × 40 pc.
Repository: GSC 137950.
Fig. 3. Hystrichostrogylon digitus sp. nov., holotype, dorso-ventral
view.
Location: Rut H-11, 2435‒2445 m. Sample: cs, P39375, 1.
Optical parameters: D, 158 × 1050, Q48/1, × 40 pc.
Repository: GSC 137952.
Fig. 4. Hystrichosphaeropsis perforata, dorso-ventral view.
Location: Skolp E-07, 1250 m. Sample: cs, YD17614, 4.
Optical parameters: A, 43.3 × 101.4, P44/1, × 60 bf.
Repository: GSC 138164.
Fig. 5. Impletosphaeridium apodastum sp. nov., holotype.
Location: Hekja O-71, 4565‒4575 m. Sample: cs, P18737, 1.
Optical parameters: C, 133 × 1080, N37/3, × 50 bf.
Repository: GSC 137903.
Fig. 6. Impletosphaeridium apodastum sp. nov.
Location: Karlsefni A-13, 3822.24–3831.38 m. Sample: cs, P39655, 1.
Optical parameters: D, 201 × 990, U42/1‒3, × 40 pc.
Repository: GSC 138055.
Fig. 7. Impletosphaeridium apodastum sp. nov.
Location: Karlsefni A-13, 3989.88–3999.02 m. Sample: cs, P39661, 1.
Optical parameters: D, 193 × 1070, T50/0, × 40 pc.
Repository: GSC 138063.
Fig. 8. Impletosphaeridium apodastum sp. nov.
Location: Karlsefni A-13, 3989.88–3999.02 m. Sample: cs, P39661, 1.
Optical parameters: D, 173 × 899, R32/0, × 40 pc.
Repository: GSC 138062.
Fig. 9. Impagidinium victorianum, ventral view of ventral surface.
Location: Roberval K-92, 2270‒2280 m. Sample: cs, P17700, 1.
Optical parameters: C, 66 × 1022, F32/3, × 50 bf.
Repository: GSC 137894.
Fig. 10. Impagidinium victorianum, ventral view, same specimen
as fig. 9, despite lower focus, ventral surface clear.
Location: Roberval K-92, 2270‒2280 m. Sample: cs, P17700, 1.
Optical parameters: C, 66 × 1022, F32/3, × 50 bf.
Repository: GSC 137894.
Fig. 11. Impagidinium victorianum, ventral view, same specimen
as fig. 9, focussed on periphery.
Location: Roberval K-92, 2270‒2280 m. Sample: cs, P17700, 1.
Optical parameters: C, 66 × 1022, F32/3, × 50 bf.
Repository: GSC 137894.
Fig. 12. Hystrichosphaeropsis quasicribrata, dorso-ventral view.
Location: North Leif I-05, 2550‒2560 m. Sample: cs, YD17614, 4.
Optical parameters: E, 43.9 × 15.8, M43/2, × 60 bf.
Repository: GSC 138165.
Fig. 13. Impagidinium victorianum, dorsal view of ventral surface.
Location: Skolp E-07, 1175 m. Sample: cs, YD15587, 3.
Optical parameters: A, 34.2 × 99.0, S435/1, × 60 bf.
Repository: MGUH 31302.
Fig. 14. Impagidinium victorianum, dorsal view of dorsal surface,
same specimen as fig. 13.
Location: Skolp E-07, 1175 m. Sample: cs, YD15587, 3.
Optical parameters: A, 34.2 × 99.0, S435/1, × 60 bf.
Repository: MGUH 31302.
Fig. 15. Isabelidinium cretaceum, dorso-ventral view.
Location: North Leif I-05, 2550‒2560 m. Sample: cs, YD17614, 2.
Optical parameters: E, 17.4 × 20.3, G15/0, × 60 bf.
Repository: MGUH 31303.
Fig. 16. Isabelidinium cretaceum, dorso-ventral view.
Location: Roberval K-92, 3060‒3070 m. Sample: cs, P17726, 1.
Optical parameters: C, 157 × 1085, Q38/2, × 50 bf.
Repository: GSC 137899.
Fig. 17. Isabelidinium microarmum, dorsal view of dorsal surface.
Location: Skolp E-07, 2285 m. Sample: cs, YD15659, 2.
Optical parameters: A, 24.4 × 93.5, X24/2, × 60 bf.
Repository: MGUH 31304.
Fig. 18. Kleithriasphaeridium mantellii, right lateral view.
Location: Maude Bight, Bylot Island, section 3 (V002181), 1 m above
base (sample PB-57). Sample: os, P5189-46, H.
Optical parameters: C, 161 × 932, L54/1–3, × 50 bf.
Repository: GSC 138131.
Fig. 19. Laciniadinium arcticum, ventral view of ventral surface.
Location: Ogmund E-72, 1530 m (reworked specimen). Sample: cs,
YD15734, 3.
Optical parameters: A, 23.6 × 98.0, S23/4, × 60 bf.
Repository: MGUH 31305.
Fig. 20. Laciniadinium arcticum, dorsal view of ventral surface.
Location: Skolp E-07, 1940 m. Sample: cs, JEH15635, 3.
Optical parameters: A, 19.5 × 104.7, L19/3, × 60 bf.
Repository: MGUH 31306.
Plate 9
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Plate 9
Plate 9
Bulletin36.qxp_Bulletin 36 19/12/16 13.40 Side 121
122
Fig. 1. Lingulodinium funginum.
Location: Karlsefni A-13, 2670.08–2679.22 m. Sample: cs, P39614, 1.
Optical parameters: D, 145 × 1047, O48/3, × 40 pc.
Repository: GSC 138042.
Fig. 2. Lingulodinium funginum.
Location: Rut H-11, 3035‒3045 m. Sample: cs, P39395, 1
Optical parameters: D, 73 × 929, G36/1, × 40 pc.
Repository: GSC 137961.
Fig. 3. Lingulodinium machaerophorum.
Location: Rut H-11, 2585‒2595 m. Sample: cs, P39380, 1.
Optical parameters: D, 105 × 900, K32/2, × 40 pc.
Repository: GSC 137954.
Fig. 4. Lingulodinium machaerophorum, dorso-ventral view show-
ing attached operculum.
Location: Gjoa O-37, 1620 m. Sample: cs, YD16074, 2.
Optical parameters: E, 18.0 × 19.6, H16/1, × 60 bf.
Repository: MGUH 31307.
Fig. 5. Nyktericysta davisii, dorso-ventral view.
Location: North Leif I-05, 3180‒3190 m. Sample: cs, YD17635, 4.
Optical parameters: E, 37.5 × 4.7, X36/0, × 40 bf.
Repository: GSC 138169.
Fig. 6. Nyktericysta davisii, dorso-ventral view.
Location: North Leif I-05, 3180‒3190 m. Sample: cs, YD17635, 4.
Optical parameters: E, 20.9 × 12.4, P19/3, × 60 bf.
Repository: GSC 138168.
Fig. 7. Nyktericysta dictyophora, dorso-ventral view.
Location: Ogmund E-72, 1995 m. Sample: cs, YD15765, 3.
Optical parameters: A, 37.3 × 94.1, W38/3, × 60 bf.
Repository: MGUH 31308.
Fig. 8. Nyktericysta dictyophora, dorso-ventral view.
Location: Ogmund E-72, 1965 m. Sample: cs, YD15763, 3.
Optical parameters: A, 23.8 × 100.8, P23/4, × 60 bf.
Repository: MGUH 31309.
Fig. 9. Lentinia serrata, dorsal view of dorsal surface.
Location: South Labrador N-79, 2700‒2710 m. Sample: cs, P39807, 1.
Optical parameters: C, 65 × 1027, F32/0–4, × 50 bf.
Repository: GSC 138122.
Fig. 10. Palaeocystodinium golzowense, dorso-ventral view.
Location: Rut H-11, 755‒765 m. Sample: cs, P39319, 1.
Optical parameters: D, 61 × 1040, E47/3, × 40 pc.
Repository: GSC 137920.
Fig. 11. Nyktericysta tripenta, dorso-ventral view; endophragm
visible apically.
Location: Ogmund E-72, 2550 m. Sample: cs, JEH15891, 3.
Optical parameters: A, 47.9 × 105.8, K49/3, × 60 bf.
Repository: MGUH 31310.
Fig. 12. Nyktericysta tripenta, dorso-ventral view, same specimen
as fig. 11.
Location: Ogmund E-72, 2550 m. Sample: cs, JEH15891, 3.
Optical parameters: A, 47.9 × 105.8, K49/3, × 60 bf.
Repository: MGUH 31310.
Fig. 13. Odontochitina porifera, dorsal view.
Location: Skolp E-07, 1895 m. Sample: cs, JEH15633, 4.
Optical parameters: A, 33.2 × 107.7, H33/4, × 60 bf.
Repository: MGUH 31311.
Fig. 14. Odontochitina ancala, dorsal view.
Location: North Leif I-05, 2790‒2800 m. Sample: cs, YD17622, 3.
Optical parameters: E, 20.1 × 18.5, J18/1, × 40 bf.
Repository: MGUH 31312.
Fig. 15. Odontochitina costata, dorsal view.
Location: Skolp E-07, 2345 m. Sample: cs, YD15663, 2.
Optical parameters: A, 40.3 × 111.8, D40/2, × 60 bf.
Repository: MGUH 31313.
Fig. 16. Licracysta corymbus, dorsal surface.
Location: Rut H-11, 1205‒1215 m. Sample: cs, P39334, 1.
Optical parameters: D, 150 × 974, P40/1‒2, × 40 pc.
Repository: GSC 137931.
Fig. 17. Oligosphaeridium albertense.
Location: Rut H-11, 1775‒1785 m. Sample: cs, P39353, 1.
Optical parameters: D, 85 × 866, H29/0, × 40 pc.
Repository: GSC 137939.
Fig. 18. Oligosphaeridium pulcherrimum, dorso-ventral view.
Location: Gilbert F-53, 2705‒2715 m. Sample: cs, P39503, 1.
Optical parameters: C, 24 × 1012, Y31/3, × 50 bf.
Repository: GSC 137986.
Fig. 19. Oligosphaeridium pulcherrimum.
Location: Rut H-11, 1745‒1755 m. Sample: cs, P39352, 1.
Optical parameters: D, 168 × 880, Q30/4, × 40 pc.
Repository: GSC 137938.
Fig. 20. Oligosphaeridium totum.
Location: Rut H-11, 2225‒2235 m. Sample: cs, P39368, 1.
Optical parameters: D, 154 × 1052, P48/0, × 40 pc.
Repository: GSC 137944.
Plate 10
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Plate 10
Plate 10
Bulletin36.qxp_Bulletin 36 19/12/16 13.40 Side 123
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Fig. 1. Palaeocystodinium bulliforme, right lateral view.
Location: Gilbert F-53, 2470‒2480 m. Sample: cs, P19228, 1.
Optical parameters: C, 163 × 1082, Q28/3, × 50 bf.
Repository: GSC 137904.
Fig. 2. Palaeocystodinium bulliforme.
Location: Ogmund E-72, 1545 m. Sample: cs, YD15735, 4.
Optical parameters: A, 30.6 × 110.6, E31/3, × 40 bf.
Repository: CNLOPB.
Fig. 3. Palaeocystodinium teespinosum, left lateral view.
Location: Karlsefni A-13, 969.28–978.42 m. Sample: cs, P39553, 1.
Optical parameters: D, 201 × 922, U35/1, × 40 pc.
Repository: GSC 138007.
Fig. 4. Palaeohystrichophora infusorioides, dorsal view.
Location: North Leif I-05, 3000‒3010 m. Sample: cs, YD17629, 3.
Optical parameters: E, 25.0 × 20.2, G23/0, × 60 bf.
Repository: MGUH 31314.
Fig. 5. Operculodinium centrocarpum.
Location: Karlsefni A-13, 694.95‒704.10 m. Sample: cs, P39543, 1.
Optical parameters: D, 143 × 996, O42/0, × 40 pc.
Repository: GSC 137996.
Fig. 6. Operculodinium centrocarpum, operculum.
Location: Karlsefni A-13, 722.38‒731.53 m. Sample: cs, P39544, 1.
Optical parameters: D, 169 × 934, R36/1‒2, × 40 pc.
Repository: GSC 137998.
Fig. 7. Palaeoperidinium pyrophorum, dorsal surface.
Location: Gilbert F-53, 2680‒2690 m. Sample: cs, P39502, 1.
Optical parameters: C, 68 × 1101, F40/3‒G40/1, × 50 bf.
Repository: GSC 137985.
Fig. 8. Palaeoperidinium pyrophorum, ventral view.
Location: Roberval K-92, 3070‒3080 m. Sample: cs, P2008175, 1.
Optical parameters: C, 200 × 982, U27/4‒U28/3, × 50 bf.
Repository: GSC 137907.
Fig. 9. Palynodinium grallator, apical view.
Location: Roberval K-92, 2840‒2850 m. Sample: cs, P17719, 1.
Optical parameters: C, 30 × 1087, C38/2–C39/1, × 50 bf.
Repository: GSC 137898.
Fig. 10. Palynodinium grallator, dorsal view.
Location: North Leif I-05, 2520‒2530 m. Sample: cs, YD17613, 4.
Optical parameters: E, 44.0 × 16.5, L46/3, × 60 bf.
Repository: GSC 138163.
Fig. 11. Petalodinium condylos, ventral view of dorsal surface.
Location: Roberval K-92, 2420‒2430 m. Sample: cs, P17705, 1.
Optical parameters: C, 110 × 1018, Q31/2–4, × 50 bf.
Repository: GSC 137896.
Fig. 12. Petalodinium condylos, ventral view of dorsal surface.
Location: Roberval K-92, 2240‒2250 m. Sample: cs, P17699, 1.
Optical parameters: C, 175 × 1052, S35/1, × 50 bf.
Repository: GSC 137892.
Fig. 13. Phthanoperidinium coreoides, dorsal surface.
Location: Rut H-11, 1145‒1155 m. Sample: cs, P39332, 1.
Optical parameters: D, 126 × 991, M42/0, × 40 pc.
Repository: GSC 137928.
Fig. 14. Phthanoperidinium coreoides, dorsal surface.
Location: North Leif I-05, 720‒730 m. Sample: cs, YD17553, 2.
Optical parameters: E, 46.7 × 11.2, Q46/1, × 60 bf.
Repository: MGUH 31315.
Fig. 15. Phthanoperidinium levimurum.
Location: Karlsefni A-13, 3273.59–3282.74 m. Sample: cs, P39636, 1.
Optical parameters: D, 100 × 1064, J50/3, × 40 pc.
Repository: GSC 138050.
Fig. 16. Phthanoperidinium multispinum, dorsal surface.
Location: South Labrador N-79, 1260‒1270 m. Sample: cs, P39762, 1.
Optical parameters: C, 87 × 1017, J31/2, × 50 bf.
Repository: GSC 138110.
Fig. 17. Phthanoperidinium multispinum, ventral surface.
Location: Karlsefni A-13, 1956.84–1965.98 m. Sample: cs, P39588, 1.
Optical parameters: D, 194 × 1025, T45/4, × 40 pc.
Repository: GSC 138020.
Fig. 18. Phthanoperidinium regale, ventral view of ventral surface.
Location: Hellefisk-1, 1289 m. Sample: cs, 02B2097-4, 4.
Optical parameters: A, 48.5 × 105.9, K49/4, × 40 bf.
Repository: MGUH 26504.
Fig. 19. Phthanoperidinium regale, ventral view of dorsal surface,
same specimen as fig. 18.
Location: Hellefisk-1, 1289 m. Sample: cs, 02B2097-4, 4.
Optical parameters: A, 48.5 × 105.9, K49/4, × 60 bf.
Repository: MGUH 26504.
Fig. 20. Phthanoperidinium stockmansii, ventral view of ventral
surface.
Location: South Labrador N-79, 1230‒1240 m. Sample: cs, P39761, 1.
Optical parameters: C, 67 × 1021, F32/3, × 50 bf.
Repository: GSC 138109.
Plate 11
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Plate 11
Plate 11
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Fig. 1. Phelodinium kozlowskii, dorsal surface.
Location: Gilbert F-53, 2740‒2750 m. Sample: cs, P39504, 1.
Optical parameters: C, 203 × 1020, U31/4–U32/3, × 50 bf.
Repository: GSC 137987.
Fig. 2. Phelodinium kozlowskii, oblique left lateral view.
Location: Bjarni O-82, 2175‒2185 m. Sample: cs, P39727, 1.
Optical parameters: C, 139 × 993, O29/1, × 50 bf.
Repository: GSC 138072.
Fig. 3. Phelodinium kozlowskii, dorsal view showing dorsal and
ventral surfaces.
Location: South Labrador N-79, 3030‒3040 m. Sample: cs, P39818, 1.
Optical parameters: C, 218 × 1000, N29/2, × 50 bf.
Repository: GSC 138123.
Fig. 4. Pseudoceratium sp., ventral view.
Location: Roberval K-92, 3070‒3080 m. Sample: cs, P2008175, 1.
Optical parameters: C, 210 × 993, V29/0‒3, × 40 pc.
Repository: GSC 137908.
Fig. 5. Piladinium columna, ventral view of dorsal surface.
Location: Bjarni O-82, 1785‒1795 m. Sample: cs, P39714, 1.
Optical parameters: C, 221 × 1054, W32/3–4, × 50 bf.
Repository: GSC 138069.
Fig. 6. Piladinium columna, dorsal view of dorsal surface.
Location: Roberval K-92, 2270‒2280 m. Sample: cs, P17700, 1.
Optical parameters: C, 200 × 1014, U31/0, × 50 bf.
Repository: GSC 137893.
Fig. 7. Piladinium edwardsii, dorsal view of dorsal surface.
Location: Ogmund E-72, 1305 m. Sample: sw, JEH15551, 4.
Optical parameters: A, 27.3 × 98.6, T27/1, × 60 bf.
Repository: MGUH 31316.
Fig. 8. Piladinium edwardsii, dorsal view of dorsal surface.
Location: North Leif I-05, 2010‒2020 m. Sample: cs, YD17596, 3.
Optical parameters: E, 21.0 × 10.6, R18/1, × 60 bf.
Repository: MGUH 31317.
Fig. 9. Raphidodinium fucatum, dorsal surface.
Location: Bjarni O-82, 2415‒2425 m. Sample: cs, P39735, 1.
Optical parameters: C, 228 × 1077, X32/0–2, × 50 bf.
Repository: GSC 138074.
Fig. 10. Raphidodinium fucatum.
Location: Skolp E-07, 1475 m. Sample: cs, YD15606, 3.
Optical parameters: A, 43.7 × 94.7, W44/1, × 60 bf.
Repository: MGUH 31318.
Fig. 11. Reticulatosphaera actinocoronata.
Location: Karlsefni A-13, 722.38‒731.53 m. Sample: cs, P39544, 1.
Optical parameters: D, 146 × 930, O36/3, × 40 pc.
Repository: GSC 137997.
Fig. 12. Reticulatosphaera actinocoronata.
Location: Rut H-11, 755‒765 m. Sample: cs, P39319, 1.
Optical parameters: D, 89 × 1006, H43/4, × 40 pc.
Repository: GSC 137921.
Fig. 13. Rhombodinium draco, ventral view of dorsal surface.
Location: Snorri J-90, 1673.37‒1682.52 m. Sample: cs, P9729, 10.
Optical parameters: C, 93 × 1007, J29/0, × 40 pc.
Repository: GSC 138134.
Fig. 14. Rhombodinium draco, ventral view of ventral surface.
Location: Bjarni O-82, 615‒625 m. Sample: cs, P39675, 1.
Optical parameters: C, 148 × 1035, P33/0, × 40 bf.
Repository: GSC 138067.
Fig. 15. Scalenodinium scalenum sp. nov., holotype, left lateral
view.
Location: Gilbert F-53, 2135‒2145 m. Sample: cs, P38484, 1.
Optical parameters: C, 193 × 1065, T37/4, × 50 bf.
Repository: GSC 137916.
Fig. 16. Scalenodinium scalenum sp. nov., left lateral view.
Location: Gilbert F-53, 2135‒2145 m. Sample: cs, P39484, 1.
Optical parameters: C, 48 × 1020, D32/3–D31/4, × 50 bf.
Repository: GSC 137982.
Fig. 17. Scalenodinium scalenum sp. nov.
Location: Ogmund E-72, 1391 m. Sample: sw, YD15553, 2.
Optical parameters: A, 26.8 × 107.0, J26/4, × 60 bf.
Repository: MGUH 31319.
Fig. 18. Scalenodinium scalenum sp. nov.
Location: Ralegh N-18, 3005 m. Sample: cs, YD16263, 4.
Optical parameters: E, 41.6 × 11.1, Q40/4, × 60 bf.
Repository: MGUH 31320.
Fig. 19. Scalenodinium scalenum sp. nov., oblique left lateral view.
Location: Ralegh N-18, 3045 m. Sample: cs, YD16264, 3.
Optical parameters: E, 28.9 × 19.9, G27/0, × 60 bf.
Repository: MGUH 31321.
Fig. 20. Scalenodinium scalenum sp. nov.
Location: North Leif I-05, 2130‒2140 m. Sample: cs, YD17600, 3.
Optical parameters: E, 29.4 × 7.6, U27/2, × 60 bf.
Repository: GSC 138156.
Plate 12
Bulletin36.qxp_Bulletin 36 19/12/16 13.40 Side 126
127
1
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4
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16
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Plate 12
Plate 12
Bulletin36.qxp_Bulletin 36 19/12/16 13.40 Side 127
128
Fig. 1. Rottnestia borussica, right lateral view.
Location: South Labrador N-79, 2430‒2440 m. Sample: cs, P39799, 1.
Optical parameters: C, 105 × 990, K28/4, × 50 bf.
Repository: GSC 138121.
Fig. 2. Schematophora speciosa, dorsal surface.
Location: South Labrador N-79, 1290‒1300 m. Sample: cs, P39763, 1.
Optical parameters: C, 126 × 982, N28/1–N27/2, × 50 bf.
Repository: GSC 138111.
Fig. 3. Senegalinium iterlaaense, dorsal surface.
Location: Gjoa O-37, 3560 m. Sample: cs, YD16131, 5.
Optical parameters: E, 46.4 × 11.6, Q45/4, × 60 bf.
Repository: GSC 138146.
Fig. 4. Senegalinium iterlaaense, ventral view of dorsal surface.
Location: Gjoa O-37, 3610 m. Sample: cs, YD16132, 3.
Optical parameters: E, 37.1 × 3.9, X36/3, × 60 bf.
Repository: MGUH 31322.
Fig. 5. Senoniasphaera inornata, dorsal view.
Location: North Leif I-05, 2490‒2500 m. Sample: cs, YD17612, 3.
Optical parameters: E, 41.2 × 8.3, T40/2, × 60 bf.
Repository: GSC 138162.
Fig. 6. Senoniasphaera microreticulata, dorsal view.
Location: North Leif I-05, 2820‒2830 m. Sample: cs, YD17623, 2.
Optical parameters: E, 41.5 × 9.2, S40/2, × 60 bf.
Repository: MGUH 31323.
Fig. 7. Senoniasphaera rotundata, dorsal view of dorsal surface.
Location: South Labrador N-79, 3480‒3490 m. Sample: cs, P39833, 1.
Optical parameters: C, 114 × 1037, L33/4, × 50 bf.
Repository: GSC 138126.
Fig. 8. Sophismatia tenuivirgula, dorso-ventral view.
Location: North Leif I-05, 1560‒1570 m. Sample: cs, YD17581, 2.
Optical parameters: E, 49.0 × 14.2, N48/2, × 60 bf.
Repository: MGUH 31324.
Fig. 9. Simplicidinium insolitum.
Location: Rut H-11, 1205‒1215 m. Sample: cs, P39334, 1.
Optical parameters: D, 152 × 871, P29/2, × 40 pc.
Repository: GSC 137932.
Fig. 10. Simplicidinium insolitum.
Location: Karlsefni A-13, 804.68–813.83 m. Sample: cs, P39547, 1.
Optical parameters: D, 162 × 1052, Q48/0, × 40 pc.
Repository: GSC 137999.
Fig. 11. Spinidinium echinoideum, dorsal view of ventral surface.
Location: Ogmund E-72, 1605 m. Sample: cs, YD15739, 3.
Optical parameters: A, 37.3 × 108.0, H38/1, × 60 bf.
Repository: MGUH 31325.
Fig. 12. Spinidinium echinoideum, dorsal view, same specimen as
fig. 11, focussed on ambitus.
Location: Ogmund E-72, 1605 m. Sample: cs, YD15739, 3.
Optical parameters: A, 37.3 × 108.0, H38/1, × 60 bf.
Repository: MGUH 31325.
Fig. 13. Spiniferites ovatus, right lateral view.
Location: Rut H-11, 755‒765 m. Sample: cs, P39319, 1.
Optical parameters: D, 135 × 933, N36/0, × 40 pc.
Repository: GSC 137923.
Fig. 14. Spiniferites pseudofurcatus.
Location: Rut H-11, 2285‒2295 m. Sample: cs, P39370, 1.
Optical parameters: D, 128 × 956, M38/4, × 40 pc.
Repository: GSC 137947.
Fig. 15. Spiniferites scabrosus.
Location: Roberval K-92, 3010 m. Sample: sw, P2008173, 1.
Optical parameters: C, 210 × 1088, V38/3–4, × 50 bf.
Repository: GSC 137906.
Fig. 16. Spongodinium delitiense, apical view.
Location: Skolp E-07, 1070 m. Sample: cs, YD15580, 4.
Optical parameters: A, 46.1× 96.5, U47/1, × 60 bf.
Repository: GSC 138143.
Fig. 17. Stichodinium lineidentatum, dorsal view of ventral surface.
Location: Nukik-2, 1862 m. Sample: cs, 05B2163, 3.
Optical parameters: A, 43.6 × 111.4, D44/4, × 60 bf.
Repository: MGUH 31326.
Fig. 18. Stichodinium lineidentatum, dorsal view of dorsal surface,
same specimen as fig. 17.
Location: Nukik-2, 1862 m. Sample: cs, 05B2163, 3.
Optical parameters: A, 43.6 × 111.4, D44/4, × 60 bf.
Repository: MGUH 31326.
Fig. 19. Spongodinium obscurum, dorsal surface.
Location: GGU 400712 borehole, 62.13‒62.25 m. Sample: cc, D309-F, 3.
Optical parameters: A, 32.0 × 109.0, G32/3, × 60 bf.
Repository: MGUH 31327.
Fig. 20. Spongodinium delitiense, right lateral view.
Location: North Leif I-05, 2460‒2470 m. Sample: cs, YD17611, 3.
Optical parameters: E, 29.2 × 18.0, J27/2, × 60 bf.
Repository: MGUH 31328.
Plate 13
Bulletin36.qxp_Bulletin 36 19/12/16 13.40 Side 128
129
32 4
17
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Plate 13
Plate 13
Bulletin36.qxp_Bulletin 36 19/12/16 13.40 Side 129
130
Fig. 1. Spongodinium grossum, oblique left lateral view.
Location: Skolp E-07, 1565 m. Sample: cs, JEH15612, 3.
Optical parameters: A, 29.5 × 110.6, E29/4, × 60 bf.
Repository: MGUH 31329.
Fig. 2. Spongodinium grossum, left lateral view.
Location: Skolp E-07, 1565 m. Sample: cs, JEH15612, 5.
Optical parameters: A, 27.3 × 98.5, S27/2, × 60 bf.
Repository: MGUH 31330.
Fig. 3. Spongodinium grossum, ventral surface.
Location: Bjarni O-82, 2445‒2455 m. Sample: cs, P39736, 1.
Optical parameters: C, 181 × 991, S29/3, × 40 bf.
Repository: GSC 138075.
Fig. 4. Spongodinium grossum, same specimen as fig. 3,
focussed on dorsal surface.
Location: Bjarni O-82, 2445‒2455 m. Sample: cs, P39736, 1.
Optical parameters: C, 181 × 991, S29/3, × 40 bf.
Repository: GSC 138075.
Fig. 5. Subtilisphaera perlucida, dorsal surface.
Location: North Leif I-05, 2910‒2920 m. Sample: cs, YD17626, 4.
Optical parameters: E, 23.4 × 19.2, H21/0, × 60 bf.
Repository: GSC 138167.
Fig. 6. Surculosphaeridium convocatum sp. nov., ventral surface.
Location: South Labrador N-79, 3510‒3520 m. Sample: cs, P39834, 1.
Optical parameters: C, 148 × 982, P28/1, × 50 bf.
Repository: GSC 138127.
Fig. 7. Surculosphaeridium convocatum sp. nov., holotype, dorsal
surface.
Location: South Labrador N-79, 3510‒3520 m. Sample: cs, P39834, 1.
Optical parameters: C, 78 × 991, G29/3–H28/2, × 50 bf.
Repository: GSC 138128.
Fig. 8. Surculosphaeridium convocatum sp. nov., lateral view.
Location: North Leif I-05, 2760‒2775 m. Sample: cs, YD17611, 2.
Optical parameters: E, 21.9 × 19.4, H20/1, × 60 bf.
Repository: MGUH 31331.
Fig. 9. Talladinium? clathratum, dorsal view, focussed on ambitus.
Location: Ralegh N-18, 1445 m. Sample: cs, YD16217, 2.
Optical parameters: E, 37.6 × 14.5, M36/4, × 40 bf.
Repository: MGUH 31332.
Fig. 10. Talladinium? clathratum, dorsal view, same specimen
as fig. 9.
Location: Ralegh N-18, 1445 m. Sample: cs, YD16217, 2.
Optical parameters: E, 37.6 × 14.5, M36/4, × 60 bf.
Repository: MGUH 31332.
Fig. 11. Talladinium pellis sp. nov., holotype, dorsal view of dorsal
surface.
Location: Gjoa O-37, 1840 m. Sample: cs, YD16082, 3.
Optical parameters: C, 21.9 × 22.0, E20/3, × 60 bf.
Repository: GSC 138082.
Fig. 12. Talladinium pellis sp. nov., holotype, dorsal view of
ventral surface, same specimen as fig. 11.
Location: Gjoa O-37, 1840 m. Sample: cs, YD16082, 3.
Optical parameters: C, 21.9 × 22.0, E20/3, × 60 bf.
Repository: GSC 138082.
Fig. 13. Taurodinium granulatum sp. nov., holotype, dorso-ventral
view.
Location: Ikermiut-1, 2340 m. Sample: cs, 2289 s-261, 3.
Optical parameters: B, 17.82 × 9.01, T19/1, × 60 bf.
Repository: MGUH 31333.
Fig. 14. Taurodinium granulatum sp. nov., dorso-ventral view.
Location: Ralegh N-18, 3045 m. Sample: cs, YD16264, 3.
Optical parameters: E, 23.6 × 14.8, M21/4, × 60 bf.
Repository: MGUH 31334.
Fig. 15. Taurodinium granulatum sp. nov., dorso-ventral view.
Location: Ralegh N-18, 3405 m. Sample: cs, YD16341, 3.
Optical parameters: E, 20.8 × 16.9, K19/3, × 60 bf.
Repository: MGUH 31335.
Fig. 16. Talladinium pellis sp. nov., ventral view.
Location: Gjoa O-37, 1840 m. Sample: cs, YD16082, 4.
Optical parameters: C, 18.56 × 15.03, L20/1, × 60 bf.
Repository: MGUH 31336.
Fig. 17. Taurodinium granulatum sp. nov., dorso-ventral view.
Location: Ralegh N-18, 3425 m. Sample: cs, YD16342, 4.
Optical parameters: E, 33.0 × 18.6, J31/2, × 60 bf.
Repository: GSC 138147.
Fig. 18. Taurodinium granulatum sp. nov.
Location: Hekja O-71, 3120 m. Sample: cs, JEH16030, 2.
Optical parameters: E, 24.9 × 23.6, C23/3, × 60 bf.
Repository: MGUH 31337.
Fig. 19. Tenua hystrix, ventral view of dorsal surface.
Location: Roberval K-92, 3090‒3100 m. Sample: cs, P17727, 1.
Optical parameters: C, 175 × 1086, S38/2, × 50 bf.
Repository: GSC 137900.
Fig. 20. Tenua hystrix, lateral view.
Location: Two Snout Creek, Bylot Island, section 538080E,
105 m above base (sample HFB-09-42). Sample: os, P514833, D.
Optical parameters: C, 145 × 918, M55/3–4, × 50 bf.
Repository: GSC 138129.
Plate 14
Bulletin36.qxp_Bulletin 36 19/12/16 13.40 Side 130
131
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Plate 14
Plate 14
Bulletin36.qxp_Bulletin 36 19/12/16 13.40 Side 131
132
Fig. 1. Thalassiphora delicata, dorsal surface.
Location: North Leif I-05, 2370‒2380 m. Sample: cs, YD17608, 4.
Optical parameters: E, 35.7 × 12.9, P34/1, × 60 bf.
Repository: MGUH 31338.
Fig. 2. Thalassiphora fenestrata, right lateral view.
Location: Ralegh N-18, 1445 m. Sample: cs, YD16217, 2.
Optical parameters: E, 32.5 × 20.5, F31/4, × 60 bf.
Repository: MGUH 31339.
Fig. 3. Thalassiphora fenestrata, left lateral view.
Location: Ralegh N-18, 1445 m. Sample: cs, YD16217, 2.
Optical parameters: E, 41.1 × 11.2, Q40/2, × 60 bf.
Repository: MGUH 31340.
Fig. 4. Thalassiphora fenestrata, dorso-ventral view.
Location: Ralegh N-18, 1605 m. Sample: cs, YD16221, 4.
Optical parameters: E, 32.8 × 6.2, V31/3, × 40 bf.
Repository: CNSOPB.
Fig. 5. Thalassiphora pelagica, dorso-ventral view.
Location: Rut H-11, 2765‒2775 m. Sample: cs, P39386, 1.
Optical parameters: D, 181 × 906, S33/0, × 40 pc.
Repository: GSC 137956.
Fig. 6. Thalassiphora pelagica, dorsal surface.
Location: Karlsefni A-13, 2615.22–2624.36 m. Sample: cs, P39612, 1.
Optical parameters: D, 52 × 902, D33/3, × 40 pc.
Repository: GSC 138039.
Fig. 7. Tanyosphaeridium xanthiopyxides.
Location: Gilbert F-53, 2590‒2600 m. Sample: cs, P39499, 1.
Optical parameters: C, 94 × 1004, J30/0, × 50 bf.
Repository: GSC 137984.
Fig. 8. Trichodinium castanea, oblique left lateral view.
Location: Skolp E-07, 1430 m. Sample: cs, YD15603, 3.
Optical parameters: A, 36.0 × 100.7, Q356/2, × 60 bf.
Repository: MGUH 31341.
Fig. 9. Trithyrodinium? conservatum sp. nov., dorso-ventral view,
with focus on ambitus.
Location: Ikermiut-1, 1155 m. Sample: sw, 04E006504, 3.
Optical parameters: A, 33.0 × 112.8, C33/3, × 60 bf.
Repository: MGUH 26500.
Fig. 10. Trithyrodinium? conservatum sp. nov., dorso-ventral view.
Location: Ikermiut-1, 1155 m. Sample: sw, 04E006504, 2.
Optical parameters: A, 41.0 × 113.5, C41/2, × 60 bf.
Repository: MGUH 26501.
Fig. 11. Trithyrodinium? conservatum sp. nov., dorso-ventral view,
same specimen as fig. 10, focussed on dorsal surface.
Location: Ikermiut-1, 1155 m. Sample: sw, 04E006504, 2.
Optical parameters: A, 41.0 × 113.5, C41/2, × 60 bf.
Repository: MGUH 26501.
Fig. 12. Trithyrodinium? conservatum sp. nov., holotype, dorsal
surface.
Location: Ikermiut-1, 1155 m. Sample: sw, 04E006504, 2.
Optical parameters: A, 36.0 × 97.1, U36/1, × 60 bf.
Repository: MGUH 26502.
Fig. 13. Trithyrodinium? conservatum sp. nov., dorsal surface.
Location: Ralegh N-18, 1485 m. Sample: cs, YD16218, 2.
Optical parameters: E, 30.3 × 23.1, D29/1, × 60 bf.
Repository: MGUH 31342.
Fig. 14. Trithyrodinium? conservatum sp. nov., dorsal surface.
Location: Ralegh N-18, 1525 m. Sample: cs, YD16219, 2.
Optical parameters: E, 28.7 × 5.8, W27/2, × 60 bf.
Repository: MGUH 31343.
Fig. 15. Trithyrodinium suspectum, dorso-ventral view.
Location: Skolp E-07, 2375 m. Sample: cs, YD15665, 4.
Optical parameters: A, 39.0 × 107.4, H39/4, × 60 bf.
Repository: MGUH 31344.
Fig. 16. Trithyrodinium quinqueangulare, oblique dorso-
ventral view.
Location: South Labrador N-79, 3300‒3310 m. Sample: cs, P39827, 1.
Optical parameters: C, 223 × 1020, W32–X31, × 50 bf.
Repository: GSC 138125.
Fig. 17. Trithyrodinium evittii, oblique apical view.
Location: North Leif I-05, 2370–2380 m. Sample: cs, YD17608, 3.
Optical parameters: E, 24.5 × 11.0, Q23/3, × 60 bf.
Repository: MGUH 31345.
Fig. 18. Trithyrodinium evittii, dorsal surface.
Location: Gilbert F-53, 2920‒2930 m. Sample: cs, P39510, 1.
Optical parameters: C, 100 × 1067, K36/3–4, × 50 bf.
Repository: GSC 137989.
Fig. 19. Trithyrodinium evittii, apical view.
Location: Roberval K-92, 3090‒3100 m. Sample: cs, P17727, 1.
Optical parameters: C, 197 × 1095, U39/0‒2, × 50 bf.
Repository: GSC 137901.
Fig. 20. Tuberculodinium vancampoae, apical view.
Location: Karlsefni A-13, 557.79–566.93 m. Sample: cs, P39538, 1.
Optical parameters: D, 146 × 1038, P46/2, × 40 pc.
Repository: GSC 137993.
Plate 15
Bulletin36.qxp_Bulletin 36 19/12/16 13.40 Side 132
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Plate 15
Plate 15
Bulletin36.qxp_Bulletin 36 19/12/16 13.40 Side 133
134
Fig. 1. Vesperopsis longicornis, dorso-ventral view.
Location: North Leif I-05, 3360–3370 m. Sample: cs, YD17641, 4.
Optical parameters: E, 55.6 × 10.1, R55/1, × 60 bf.
Repository: MGUH 31346.
Fig. 2. Vesperopsis longicornis, dorso-ventral view.
Location: North Leif I-05, 3360–3370 m. Sample: cs, YD17641, 3.
Optical parameters: E, 35.2 × 23.4, C34/1, × 60 bf.
Repository: MGUH 31347.
Fig. 3. Wetzeliella articulata, dorsal view.
Location: Rut H-11, 3725‒3735 m. Sample: cs, P39417, 1.
Optical parameters: D, 152 × 847, P27/0, × 40 pc.
Repository: GSC 137971.
Fig. 4. Wetzeliella articulata, dorsal view.
Location: Rut H-11, 3095‒3105 m. Sample: cs, P39397, 1.
Optical parameters: D, 81 × 949, G38/3, × 40 pc.
Repository: GSC 137963.
Fig. 5. Wallodinium luna, lateral view.
Location: Skolp E-07, 1610 m. Sample: cs, JEH15615, 3.
Optical parameters: A, 39.6 × 100.2, R40/1, × 60 bf.
Repository: MGUH 31348.
Fig. 6. Xenascus wetzelii, dorsal view of dorsal surface.
Location: Skolp E-07, 1475 m. Sample: cs, YD15606, 3.
Optical parameters: A, 41.0 × 111.2, E41/2, × 40 bf.
Repository: MGUH 31349.
Fig. 7. Xenascus wetzelii, dorsal view, same specimen as fig. 6,
showing enlarged view of ventral surface.
Location: Skolp E-07, 1475 m. Sample: cs, YD15606, 3.
Optical parameters: A, 41.0 × 111.2, E41/2, × 60 bf.
Repository: MGUH 31349.
Fig. 8. Xenascus wetzelii, dorsal view, same specimen as fig. 6,
showing enlarged view of dorsal surface.
Location: Skolp E-07, 1475 m. Sample: cs, YD15606, 3.
Optical parameters: A, 41.0 × 111.2, E41/2, × 60 bf.
Repository: MGUH 31349.
Fig. 9. Xenascus ceratioides, ventral view.
Location: Rut H-11, 2255‒2235 m. Sample: cs, P39369, 1.
Optical parameters: D, 116 × 954, L38/0, × 40 pc.
Repository: GSC 137945.
Fig. 10. Xenascus ceratioides, dorsal view.
Location: Skolp E-07, 2435 m. Sample: cs, YD15669, 3.
Optical parameters: A, 24.6 × 107.9, H24/2, × 60 bf.
Repository: GSC 138144.
Fig. 11. Fromea quadrangularis sp. nov.
Location: Skolp E-07, 2075 m. Sample: cs, JEH15643, 3.
Optical parameters: A, 42.1 × 94.3, W43/3, × 60 bf.
Repository: MGUH 31350.
Fig. 12. Fromea quadrangularis sp. nov.
Location: Skolp E-07, 2090 m. Sample: cs, JEH15644, 5.
Optical parameters: A, 44.2 × 109.3, G45/1, × 60 bf.
Repository: GSC 137882.
Fig. 13. Microsphaeridium ancistroides.
Location: Rut H-11, 665‒675 m. Sample: cs, P39316, 1.
Optical parameters: D, 108 × 900, K33/3, × 40 pc.
Repository: GSC 137917.
Fig. 14. Microsphaeridium ancistroides.
Location: Rut H-11, 1835‒1845 m. Sample: cs, P39355, 1.
Optical parameters: D, 186 × 907, S33/3‒4, × 40 pc.
Repository: GSC 137941.
Fig. 15. Fromea quadrangularis sp. nov.
Location: Skolp E-07, 2360 m. Sample: cs, YD15664, 2.
Optical parameters: A, 47.6 × 99.1, R45/4, × 60 bf.
Repository: MGUH 31351.
Fig. 16. Fromea quadrangularis sp. nov., holotype.
Location: Skolp E-07, 2375 m. Sample: cs, YD15665, 4.
Optical parameters: A, 30.9 × 111.9, D31/1, × 60 bf.
Repository: MGUH 31352.
Fig. 17. Microsphaeridium ancistroides.
Location: Karlsefni A-13, 667.52–676.66 m. Sample: cs, P39542, 1.
Optical parameters: D, 170 × 946, R37/2, × 40 pc.
Repository: GSC 137994.
Fig. 18. Microsphaeridium ancistroides.
Location: Rut H-11, 665‒675 m. Sample: cs, P39316, 1.
Optical parameters: D, 135 × 1003, N43/0, × 40 pc.
Repository: GSC 137918.
Fig. 19. Fromea nicosia.
Location: Skolp E-07, 2360 m. Sample: cs, YD15664, 2.
Optical parameters: A, 25.5 × 93.1, X25/0, × 60 bf.
Repository: MGUH 31353.
Fig. 20. Palambages sp.
Location: Rut H-11, 2405‒2415 m. Sample: cs, P39374, 1.
Optical parameters: D, 169 × 1003, R43/1‒2, × 40 pc.
Repository: GSC 137951.
Plate 16
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Plate 16
Plate 16
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136
Fig. 1. Paralecaniella indentata.
Location: Karlsefni A-13, 2176.30–2185.44 m. Sample: cs, P39596, 1.
Optical parameters: D, 162 × 873, Q30/1, × 40 pc.
Repository: GSC 138024.
Fig. 2. Paralecaniella indentata.
Location: Hellefisk-1, 1588 m. Sample: sw, 02E6529-3, 3.
Optical parameters: A, 30.5 × 106.3, K30/0, × 60 bf.
Repository: MGUH 26544.
Fig. 3. Pediastrum sp.
Location: Rut H-11, 1865‒1875 m. Sample: cs, P39356, 1.
Optical parameters: D, 136 × 1057, N49/3, × 40 pc.
Repository: GSC 137943.
Fig. 4. Pediastrum sp.
Location: Karlsefni A-13, 2286.03–2295.17 m. Sample: cs, P39600, 1.
Optical parameters: D, 167 × 1007, Q44/3, × 40 pc.
Repository: GSC 138032.
Fig. 5. Tetraporina sp. A.
Location: Gjoa O-37, 1680 m. Sample: cs, YD16076, 2.
Optical parameters: E, 23.1 × 14.4, N21/0, × 60 bf.
Repository: MGUH 31354.
Fig. 6. Tetraporina sp. A.
Location: North Leif I-05, 540‒550 m. Sample: cs, YD17547, 3.
Optical parameters: E, 37.0 × 11.3, Q35/4, × 60 bf.
Repository: GSC 138148.
Fig. 7. Tetraporina sp. A.
Location: Karlsefni A-13, 2011.70–2020.85 m. Sample: cs, P39590, 1.
Optical parameters: D, 191 × 923, T35/0, × 40 pc.
Repository: GSC 138021.
Fig. 8. Tetraporina sp. B.
Location: Rut H-11, 3035‒3045 m. Sample: cs, P39395, 1.
Optical parameters: D, 134 × 931, N36/0, × 40 pc.
Repository: GSC 137962.
Fig. 9. Tetraporina sp. B.
Location: Gjoa O-37, 1620 m. Sample: cs, YD16074, 4.
Optical parameters: A, 42.5 × 98.2, S43/0, × 60 bf.
Repository: MGUH 31355.
Fig. 10. Tetraporina sp. B.
Location: Gjoa O-37, 1620 m. Sample: cs, YD16074, 4.
Optical parameters: A, 34.0 × 105.0, L34/0, × 60 bf.
Repository: MGUH 31356.
Fig. 11. Tetraporina sp. B.
Location: Skolp E-07, 935 m. Sample: cs, YD15568, 4.
Optical parameters: A, 22.5 × 98.4, S22/2, × 60 bf.
Repository: GSC 138141.
Fig. 12. Tetraporina sp. B.
Location: Rut H-11, 3485‒3495 m. Sample: cs, P39410, 1.
Optical parameters: D, 164 × 904, Q33/1‒3, × 40 pc.
Repository: GSC 137968.
Fig. 13. Translucentipollis contiguus.
Location: Rut H-11, 3785‒3795 m. Sample: cs, P39419, 1.
Optical parameters: D, 155 × 978, P40/4, × 40 pc.
Repository: GSC 137973.
Fig. 14. Aquilapollenites quadrilobus.
Location: Karlsefni A-13, 1956.84–1965.98 m. Sample: cs, P39588, 1.
Optical parameters: D, 138 × 1037, N47/3, × 40 pc.
Repository: GSC 138019.
Fig. 15. Aquilapollenites quadrilobus.
Location: Bjarni O-82, 799‒800 m. Sample: cs, P39681, 1.
Optical parameters: C, 220 × 1048, W34/4, × 40 bf.
Repository: GSC 138068.
Fig. 16. Parviprojectus reticulatus.
Location: Rut H-11, 3605‒3615 m. Sample: cs, P39413, 1.
Optical parameters: D, 198 × 855, U28/1, × 40 pc.
Repository: GSC 137969.
Fig. 17. Aquilapollenites quadrilobus.
Location: Hekja O-71, 2080 m. Sample: cs, JEH16005, 1.
Optical parameters: C, 145 × 1007, O30/4–P30/2, × 50 bf.
Repository: GSC 137884.
Fig. 18. Afropollis sp.
Location: Ogmund E-72, 1845 m. Sample: cs, YD15755, 3.
Optical parameters: A, 26.4 × 101.8, O26/4, × 60 bf.
Repository: MGUH 31357.
Fig. 19. Afropollis sp.
Location: Ogmund E-72, 1845 m. Sample: cs, YD15755, 3.
Optical parameters: A, 26.8 × 96.7, U27/1, × 60 bf.
Repository: MGUH 31358.
Fig. 20. Appendicisporites potomacensis.
Location: Ogmund E-72, 2340 m. Sample: cs, JEH15877, 3.
Optical parameters: A, 16.3 × 111.3, D16/3, × 60 bf.
Repository: MGUH 31359.
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Fig. 1. Appendicisporites unicus.
Location: Ogmund E-72, 2205 m. Sample: cs, JEH15868, 3.
Optical parameters: A, 28.6 × 102.6, N28/4, × 60 bf.
Repository: GSC 137883.
Fig. 2. Azolla sp.
Location: Ralegh N-18, 2885 m. Sample: cs, YD16260, 3.
Optical parameters: E, 25.1 × 8.1, T23/2, × 60 bf.
Repository: MGUH 31360.
Fig. 3. Azolla sp.
Location: Gjoa O-37, 1858‒1868 m. Sample: cs, P16963, 1.
Optical parameters: C, 50 × 1011, E31/1, × 50 bf.
Repository: GSC 137886.
Fig. 4. Azolla sp.
Location: Karlsefni A-13, 2395.76–2404.90 m. Sample: cs, P39604, 1.
Optical parameters: D, 153 × 1034, P46/2‒4, × 40 pc.
Repository: GSC 138034.
Fig. 5. Cicatricososporites eocenicus.
Location: South Labrador N-79, 2040‒2050 m. Sample: cs, P39786, 1.
Optical parameters: C, 187 × 1040, T32/1–T33/2, × 50 bf.
Repository: GSC 138118.
Fig. 6. Baculatisporites crenulatus sp. nov.
Location: Roberval K-92, 2090‒2100 m. Sample: cs, P17692, 1.
Optical parameters: C, 45 × 1091, D39/0, × 50 bf.
Repository: GSC 137890.
Fig. 7. Baculatisporites crenulatus sp. nov.
Location: Roberval K-92, 2150‒2160 m. Sample: cs, P17696, 1.
Optical parameters: C, 63 × 1073, F37/0, × 50 bf.
Repository: GSC 137891.
Fig. 8. Baculatisporites crenulatus sp. nov., holotype.
Location: Roberval K-92, 1790‒1800 m. Sample: cs, P17684, 1.
Optical parameters: C, 106 × 1048, L34/2, × 50 bf.
Repository: GSC 137889.
Fig. 9. Caryapollenites inelegans.
Location: Karlsefni A-13, 969.28–978.42 m. Sample: cs, P39553, 1.
Optical parameters: D, 81 × 1060, H49/2, × 40 pc.
Repository: GSC 138003.
Fig. 10. Caryapollenites inelegans.
Location: Rut H-11, 845‒855 m. Sample: cs, P39322, 1.
Optical parameters: D, 112 × 844, L27/1, × 40 pc.
Repository: GSC 137925.
Fig. 11. Caryapollenites veripites.
Location: Karlsefni A-13, 2231.16–2240.31 m. Sample: cs, P39598, 1.
Optical parameters: D, 191 × 993, T42/0, × 40 pc.
Repository: GSC 138028.
Fig. 12. Caryapollenites veripites.
Location: Karlsefni A-13, 3843.57–3852.72 m. Sample: cs, P39656, 1.
Optical parameters: D, 164 × 867, Q29/0, × 40 pc.
Repository: GSC 138056.
Fig. 13. Momipites annellus.
Location: Karlsefni A-13, 3904.54–3913.68 m. Sample: cs, P39658, 1.
Optical parameters: D, 175 × 911, R34/3, × 40 pc.
Repository: GSC 138058.
Fig. 14. Momipites annellus.
Location: Karlsefni A-13, 3301.02–3310.17 m. Sample: cs, P39637, 1.
Optical parameters: D, 122 × 917, M34/0, × 40 pc.
Repository: GSC 138051.
Fig. 15. Momipites coryloides.
Location: Snorri J-90, 1033.28‒1043.43 m. Sample: cs, P9709, 10.
Optical parameters: C, 103 × 1078, K37–K38, × 50 bf.
Repository: GSC 138133.
Fig. 16. Callialasporites dampieri.
Location: Karlsefni A-13, 969.28–978.42 m. Sample: cs, P39553, 1.
Optical parameters: D, 196 × 1026, T45/4, × 40 pc.
Repository: GSC 138006.
Fig. 17. Chenopodipollis sp.
Location: Karlsefni A-13, 911.36–920.51 m. Sample: cs, P39551, 1.
Optical parameters: D, 115 × 930, L35/2‒4, × 40 pc.
Repository: GSC 138001.
Fig. 18. Compositoipollenites sp. B. of Williams & Brideaux 1975.
Location: Karlsefni A-13, 1435.63–1444.77 m. Sample: cs, P39570, 1.
Optical parameters: D, 187 × 934, S36/3, × 40 pc.
Repository: GSC 138012.
Fig. 19. Corsinipollenites oculusnoctis.
Location: South Labrador N-79, 1110‒1120 m. Sample: cs, P39757, 1.
Optical parameters: C, 200 × 1018, U31/1–4, × 50 bf.
Repository: GSC 138080.
Fig. 20. Corsinipollenites oculusnoctis.
Location: Karlsefni A-13, 1709.95–1719.09 m. Sample: cs, P39579, 1.
Optical parameters: D, 196 × 1042, T47/3, × 40 pc.
Repository: GSC 138014.
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Fig. 1. Cicatricosisporites ornatus.
Location: Rut H-11, 2315‒2325 m. Sample: cs, P39371, 1.
Optical parameters: D, 163 × 885, Q31/0, × 40 pc.
Repository: GSC 137949.
Fig. 2. Cicatricosisporites ornatus.
Location: Rut H-11, 1865‒1875 m. Sample: cs, P39356, 1.
Optical parameters: D, 115 × 881, L30/2‒4, × 40 pc.
Repository: GSC 137942.
Fig. 3. Cicatricosisporites ornatus.
Location: Karlsefni A-13, 2532.98–2542.08 m. Sample: cs, P39609, 1.
Optical parameters: D, 163 × 873, Q30/1‒3, × 40 pc.
Repository: GSC 138038.
Fig. 4. Cicatricosisporites ornatus.
Location: Rut H-11, 3695‒3705 m. Sample: cs, P39416, 1.
Optical parameters: D, 90 × 993, H42/3‒4, × 40 pc.
Repository: GSC 137970.
Fig. 5. Extratriporopollenites sp.
Location: Gilbert F-53, 1540‒1550 m. Sample: cs, P39464, 1.
Optical parameters: C, 180 × 1043, S34/0, × 50 bf.
Repository: GSC 137975.
Fig. 6. Extratriporopollenites sp.
Location: Hekja O-71, 1610‒1620 m. Sample: cs, P20568, 1.
Optical parameters: C, 197 × 1032, U33/1, × 50 bf.
Repository: GSC 137914.
Fig. 7. Extratriporopollenites sp.
Location: Hekja O-71, 1420‒1430 m. Sample: cs, P20562, 1.
Optical parameters: C, 80 × 1077, H37/2, × 50 bf.
Repository: GSC 137913.
Fig. 8. Extratriporopollenites sp.
Location: Snorri J-90, 1892.83‒1901.98 m. Sample: cs, P9737, 1.
Optical parameters: C, 135 × 1020, N31/4–N32/3, × 50 bf.
Repository: GSC 138135.
Fig. 9. Extratriporopollenites sp.
Location: South Labrador N-79, 780‒790 m. Sample: cs, P39746, 1.
Optical parameters: C, 154 × 1018, P31/4, × 50 bf.
Repository: GSC 138078.
Fig. 10. Periporopollenites sp.
Location: Karlsefni A-13, 2176.30–2185.44 m. Sample: cs, P39596, 1.
Optical parameters: D, 109 × 957, K38/4, × 40 pc.
Repository: GSC 138023.
Fig. 11. Periporopollenites sp.
Location: Rut H-11, 1305–1315 m. Sample: cs, P39337, 1.
Optical parameters: D, 148 × 859, O28/0, × 40 pc.
Repository: GSC 137935.
Fig. 12. Graminidites sp. A of Williams & Brideaux 1975.
Location: Rut H-11, 1685‒1695 m. Sample: cs, P39350, 1.
Optical parameters: D, 177 × 882, R31/3, × 40 pc.
Repository: GSC 137937.
Fig. 13. Pistillipollenites macgregorii.
Location: Snorri J-90, 2441.48‒2450.62 m. Sample: cs, P9751, 10.
Optical parameters: C, 30 × 910, C20/0, × 50 bf.
Repository: GSC 138140.
Fig. 14. Pistillipollenites macgregorii.
Location: Gjoa O-37, 1620‒1630 m. Sample: cs, P16951, 1.
Optical parameters: C, 65 × 1174, F37/4, × 50 bf.
Repository: GSC 137885.
Fig. 15. Quercoidites sp.
Location: Rut H-11, 1205‒1215 m. Sample: cs, P39334, 1.
Optical parameters: D, 119 × 869, L29/4, × 40 pc.
Repository: GSC 137930.
Fig. 16. Quercoidites sp.
Location: Rut H-11, 1305‒1315 m. Sample: cs, P39337, 1.
Optical parameters: D, 102 × 957, K38/2, × 40 pc.
Repository: GSC 137934.
Fig. 17. Tiliaepollenites crassipites.
Location: South Labrador N-79, 1110‒1120 m. Sample: cs, P39757, 1.
Optical parameters: C, 182 × 1009, S30/2, × 50 bf.
Repository: GSC 138079.
Fig. 18. Tiliaepollenites crassipites.
Location: South Labrador N-79, 780‒790 m. Sample: cs, P39746, 1.
Optical parameters: C, 134 × 1023, N32/3, × 50 bf.
Repository: GSC 138077.
Fig. 19. Tiliaepollenites sp. A.
Location: Rut H-11, 1565‒1575 m. Sample: cs, P39346, 1.
Optical parameters: D, 190 × 928, T35/2, × 40 pc.
Repository: GSC 137936.
Fig. 20. Tiliaepollenites sp. A.
Location: Karlsefni A-13, 911.36–920.51 m. Sample: cs, P39551, 1.
Optical parameters: D, 134 × 1000, N43/0, × 40 pc.
Repository: GSC 138002.
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Fig. 1. Osmundacidites wellmannii.
Location: Rut H-11, 905‒915 m. Sample: cs, P39324, 1.
Optical parameters: D, 191 × 968, T39/2, × 40 pc.
Repository: GSC 137927.
Fig. 2. Rugubivesiculites sp.
Location: Ogmund E-72, 1965 m. Sample: cs, YD15763, 3.
Optical parameters: A, 35.4 × 103.2, N35/2, × 60 bf.
Repository: MGUH 31361.
Fig. 3. Wodehouseia spinata.
Location: Skolp E-07, 1015 m. Sample: cs, YD15576, 3.
Optical parameters: A, 52.0 × 100.5, Q53/1, × 60 bf.
Repository: MGUH 31362.
Fig. 4. Zlivisporis sp.
Location: Karlsefni A-13, 1792.25–1801.39 m. Sample: cs, P39582, 1.
Optical parameters: D, 96 × 851, J27/2, × 40 pc.
Repository: GSC 138017.
Fig. 5. Zlivisporis sp.
Location: Rut H-11, 2285‒2295 m. Sample: cs, P39370, 1.
Optical parameters: D, 133 × 1041, N47/0, × 40 pc.
Repository: GSC 137948.
Fig. 6. Zlivisporis sp.
Location: Karlsefni A-13, 2148.87–2158.01 m. Sample: cs, P39595, 1.
Optical parameters: D, 110 × 1047, K48/3, × 40 pc.
Repository: GSC 138022.
Fig. 7. Zlivisporis sp.
Location: Rut H-11, 1805‒1815 m. Sample: cs, P39354, 1.
Optical parameters: D, 120 × 1031, M46/1, × 40 pc.
Repository: GSC 137940.
Fig. 8. Zlivisporis sp.
Location: Ogmund E-72, 1785 m. Sample: cs, YD15751, 3.
Optical parameters: A, 17.5 × 111.0, E17/2, × 60 bf.
Repository: MGUH 31363.
Fig. 9. Zonalapollenites igniculus.
Location: Karlsefni A-13, 859.55–865.64 m. Sample: cs, P39549, 1.
Optical parameters: D, 100 × 954, J38/3‒4, × 40 pc.
Repository: GSC 138000.
Fig. 10. Zonalapollenites igniculus.
Location: Rut H-11, 845‒855 m. Sample: cs, P39322, 1.
Optical parameters: D, 179 × 914, S34/1, × 40 pc.
Repository: GSC 137926.
Fig. 11. Zonalapollenites igniculus.
Location: South Labrador N-79, 720‒730 m. Sample: cs, P39744, 1.
Optical parameters: C, 142 × 1009, O30/4–P30/2, × 50 bf.
Repository: GSC 138076.
Fig. 12. Fungal element: Callimothallus.
Location: Karlsefni A-13, 2478.05–2487.20 m. Sample: cs, P39607, 1.
Optical parameters: D, 161 × 1054, Q48/0, × 40 pc.
Repository: GSC 138035.
Fig. 13. Fungal element: Diporicellaesporites.
Location: Karlsefni A-13, 2505.49–2514.63 m. Sample: cs, P39608, 1.
Optical parameters: D, 127 × 964, M39/0, × 40 pc.
Repository: GSC 138036.
Fig. 14. Fungal element: Fractisporonites.
Location: Karlsefni A-13, 1737.38–1746.53 m. Sample: cs, P39580, 1.
Optical parameters: D, 179 × 889, R31/4, × 40 pc.
Repository: GSC 138015.
Fig. 15. Fungal element: Fusiformisporites.
Location: Rut H-11, 2285‒2295 m. Sample: cs, P39370, 1.
Optical parameters: D, 106 × 906, K33/0, × 40 pc.
Repository: GSC 137946.
Fig. 16. Fungal element: Microthallites.
Location: Karlsefni A-13, 1764.81–1773.96 m. Sample: cs, P39581, 1.
Optical parameters: D, 108 × 887, K31/0, × 40 pc.
Repository: GSC 138016.
Fig. 17. Fungal element: Staphlosporonites.
Location: Karlsefni A-13, 2286.03–2295.17 m. Sample: cs, P39600, 1.
Optical parameters: D, 141 × 1071, O50/0, × 40 pc.
Repository: GSC 138031.
Fig. 18. Fungal element: Multicellaesporites.
Location: Karlsefni A-13, 1792.25–1801.39 m. Sample: cs, P39582, 1.
Optical parameters: D, 186 × 885, S31/0, × 40 pc.
Repository: GSC 138018.
Fig. 19. Fungal element: Multicellaesporites.
Location: Rut H-11, 3725‒3735 m. Sample: cs, P39417, 1.
Optical parameters: D, 175 × 1043, R47/4, × 40 pc.
Repository: GSC 137972.
Fig. 20. Fungal element: Pluricellaesporites.
Location: Rut H-11, 2435‒2445 m. Sample: cs, P39375, 1.
Optical parameters: D, 160 × 887, Q31/1‒2, × 40 pc.
Repository: GSC 137953.
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<<
/AddBleedMarks false
/AddColorBars false
/AddCropMarks true
/AddPageInfo false
/AddRegMarks false
/BleedOffset [
8.503940
8.503940
8.503940
8.503940
]
/ConvertColors /NoConversion
/DestinationProfileName ()
/DestinationProfileSelector /NA
/Downsample16BitImages true
/FlattenerPreset <<
/PresetSelector /HighResolution
>>
/FormElements false
/GenerateStructure false
/IncludeBookmarks false
/IncludeHyperlinks false
/IncludeInteractive false
/IncludeLayers false
/IncludeProfiles false
/MarksOffset 6
/MarksWeight 0.250000
/MultimediaHandling /UseObjectSettings
/Namespace [
(Adobe)
(CreativeSuite)
(2.0)
]
/PDFXOutputIntentProfileSelector /DocumentCMYK
/PageMarksFile /RomanDefault
/PreserveEditing true
/UntaggedCMYKHandling /LeaveUntagged
/UntaggedRGBHandling /LeaveUntagged
/UseDocumentBleed false
>>
<<
/AllowImageBreaks true
/AllowTableBreaks true
/ExpandPage false
/HonorBaseURL true
/HonorRolloverEffect false
/IgnoreHTMLPageBreaks false
/IncludeHeaderFooter false
/MarginOffset [
0
0
0
0
]
/MetadataAuthor ()
/MetadataKeywords ()
/MetadataSubject ()
/MetadataTitle ()
/MetricPageSize [
0
0
]
/MetricUnit /inch
/MobileCompatible 0
/Namespace [
(Adobe)
(GoLive)
(8.0)
]
/OpenZoomToHTMLFontSize false
/PageOrientation /Portrait
/RemoveBackground false
/ShrinkContent true
/TreatColorsAs /MainMonitorColors
/UseEmbeddedProfiles false
/UseHTMLTitleAsMetadata true
>>
]
>> setdistillerparams
<<
/HWResolution [2400 2400]
/PageSize [612.000 792.000]
>> setpagedevice