Stratigraphy and Paleoenvironmental Interpretation of the ...€¦ · Recent study of foraminifera, nummulites and nannofossils (Bugrova et al., 2002) from sandyclayey sequence revealed

ISSN 0869�5938, Stratigraphy and Geological Correlation, 2011, Vol. 19, No. 4, pp. 424–449. © Pleiades Publishing, Ltd., 2011.Original Russian Text © G.N. Aleksandrova, E.A. Shcherbinina, 2011, published in Stratigrafiya. Geologicheskaya Korrelyatsiya, 2011, Vol. 19, No. 4, pp. 62–88.

424

INTRODUCTION

One of the major paleoecological crises amongsuccessive Mesozoic�Cenozoic biotic events is so�called Paleocene–Eocene Thermal Maximum(PETM). By development rate and many otherparameters, the PETM is comparable with present cli�matic events. Recently, it has been shown that dra�matic disruption of the carbon cycle and climaticwarming associated with the PETM caused negativecarbon and oxygen isotope excursions in the geologi�cal record (Kennett and Stott, 1991; Zachos et al.,2001; a.o.), reorganization of the oceanic current sys�tem (Nunes and Norris, 2006; Tripati and Elderfield,2005), dramatic compositional turnovers in marineand terrestrial biotas (Thomas, 1998; Gibbs et al.,2006; Clyde and Gingerich, 1998; Wing et al., 2005),and widespread occurrence of sediments enriched inorganic matter (Gavrilov et al., 2003; Speijer and Wag�ner, 2002). Despite extensive study of involved intoPETM phenomena and the great volume of recentlyobtained information, the causes of its occurrence andprecise succession of the associated events are stillpoorly understood.

In the northeastern Peri�Tethys, the PETM is pro�nounced as sapropelitic bed (SB, Muzylev, 1994) fea�tured by significant enrichment in TOC, drastic car�bon and oxygen isotope excursions (up to 20%,Gavrilov et al., 1997), and prominent turnovers inmicroplankton assemblages (Stupin and Muzylev,2001; Gavrilov et al., 2003). The study of nannofossilvariations within SB of Caucasian and Central Asia

sections revealed dramatic decrease in nannofossilabundance comparing to embedding sediments,occurrence of so�called “excursion Taxa” (e.g. thespecies whose stratigraphic distribution ranges CarbonIsotope Excursion � CIE) namely Rhomboaster spp.and asymmetrical discoasterids (Discoaster anartiosand D. araneus), and changes in relative abundancesof taxa caused warm�water and eutrophic speciesdomination in the assemblages (Gavrilov et al., 2003).Besides, domination of Apectodinium genus andoccurrence of “excursion species” A. augustum areshown in some areas (Akhmetiev and Zaporozhets,1996; Iakovleva et al., 2001; Crouch et al., 2003).

Paleogene deposits in the Eastern Crimea arepoorly exposed, and thus inadequately studied. Out�crops of Paleogene rocks in unnamed gullies south�wards of the village of Nasypnoe (6 km westward of thecity of Feodosiya) are better exposed than elsewhere inthe area. This succession is the reference Paleogenesection for the Eastern Crimea (Stratigrafiya…, 1984).The succession includes, from bottom to top, theDanian Feodosiya Formation, uncomformably over�lain by sandy�clayey sequence tentatively correlatedwith the Thanetian Kacha Horizon of WesternCrimea, which is superposed by the Nasypkoi Stratacorrelative with the Eocene Bakhchisarai and Simfer�opol horizons. Despite the multiple studies (Nemkovand Barkhatova, 1961; Shutskaya, 1970; Gorbach,1972, a.o.), the age of lithostratigraphic units remainsproblematic.

Stratigraphy and Paleoenvironmental Interpretation of the Paleocene–Eocene Transition in the Eastern Crimea

G. N. Aleksandrova and E. A. ShcherbininaGeological Institute, Russian Academy of Sciences, Moscow, Russia; e�mail: [email protected]

Received March 2, 2010; in final form, April 27, 2010

Abstract—The study of nannofossils and dinoflagelate cysts from the Paleocene–Eocene transition in theNasypnoe section, Eastern Crimea identified the bed corresponding to the global event referred as the Paleo�cene–Eocene Thermal Maximum (PETM). The assemblages of both groups of microphytoplankton displaysignificant changes including the appearance of Rhomboaster spp., Discoaster anartios and D. araneus nanno�fossils and Apectodinium augustum and Wilsonidium pechoricum dinocysts featured for this event and majorvariations in the ratio of taxa resulted in domination of eutrophic and warm�water species. The paleoecolog�ical interpretation of nannofossil and dinocyst distribution suggests a drastic sea�level fall preceded thePETM and occurrence of two transgressive episodes during it.

Keywords: biostratigraphy, Paleocene–Eocene Thermal Maximum, dinocysts, nannoplankton, EasternCrimea.

DOI: 10.1134/S0869593811040022

STRATIGRAPHY AND GEOLOGICAL CORRELATION Vol. 19 No. 4 2011

STRATIGRAPHY AND PALEOENVIRONMENTAL INTERPRETATION 425

Recent study of foraminifera, nummulites andnannofossils (Bugrova et al., 2002) from sandy�clayeysequence revealed its late Selandian to Thanetian age.The age of sequence was defined as non�subdividedinterval of NP6�NP9 nannofossil zones, Acarininaacarinata planktonic foraminifera zone, Karreiellazolkaensis benthic foraminifera zone, and SBZ3 num�mulite zone (at the base of the section). The stratigra�phy of the upper part of this sequence and transition tothe Nasypkoi Strata remained unresolved, because ofthe gap in field observation. The overlying NasypkoiStrata were dated as early Ypresian and correlated withupper part of the Bakhchisarai Fm. and basal Simfer�opol beds of the stratotype region (Western Crimea).Bugrova et al. (2002) noted that microfossils fromupper part of the sandy�clayey sequence should bestudied in greater detail as this interval contains char�acteristic agglutinated foraminifers of the Karreiellazolkaensis zone which are indicators of oxygen defi�ciency in bottom waters. In the same part of the sec�tion, the appearance of planktonic foraminifers of theMorozovella aequa zone indicating the Paleocene–Eocene transition was recorded. However, the actuallevel of Paleocene–Eocene boundary has not beenrecognized because neither foraminiferal nor nanno�fossil markers were found in the interval studied. Asimilar assemblage was identified by authors in thesections of Kerchenskii Peninsula, Crimea and low�calcareous to carbonate�free deposits of the GoryachiiKlyuch Fm. in the northwestern Caucasus and centralCiscaucasia.

Our interest in this sandy�clayey sequence is basedon the possible occurrence of the beds related toPETM. Apparently, the Nasypnoe section of easternCrimea represents the westernmost region of PETMmanifestation in the northeastern Peri�Tethys, wherethis interval is composed of mostly siliciclastic sedi�ments. In westerly areas, for instance Suvlukaya sec�tion near Bakhchisarai, the Paleocene–Eocene transi�tion corresponds to hiatus spanning the NP9�NP10nannofossil zones. To reveal the peculiar microfossilchanges across the Paleocene–Eocene boundary inthe eastern Crimea, we thoroughly studied nannofossiland dinocyst assemblages and recognized biostrati�graphic zones based on both groups. Dinocysts of theNasypnoe sections were studied for the first time.

MATERIALS AND METHODS

Lithology of Studied Section

In 2008, Yu.O. Gavrilov and E.A. Shcherbininasampled the section exposed in an unnamed gully(45°02′00′′N, 35°17′11′′E) southwestward of the vil�lage of Nasypnoe. Rock samples have been collectedfrom isolated test pits in the covered intervals on thegully slopes; hence the lithological boundaries are onlyprovisionally defined. The following lithological unitshave been identified in the right slope of the gully frombase to top (Fig. 1):

Feodosiya Formation

1. Compact yellowish gray bioclastic limestone;exposed thickness is ~2 m.

2. White, highly porous, clastic Lithothamniumlimestone with fragments of mollusks and coal clasts;thickness varies along the strike, commonly ranging0.5 to 0.8 m.

Upsection, 1.8 to 2.0 m thick interval is unexposed.

Sandy�Clayey Sequence

3. Gray soft sandy to silty clay with isolated severalcm thick intercalations of bioclastic material in thelower part; thickness is ~2.2 m.

4. Soft yellowish�gray 0.6 m thick sandy clay.5. Plastic calcareous clay ~1.3 m thick, gray in the

lower part and dark gray in the upper part.6. Rusty�gray clay ~1.7 m thick with rare silty inter�

calations.7. Gray laminated mudstone; discovered thickness

~3.5 m.Hence, the sapropelitic horizon, very distinct in

the eastern sections of the northeastern Peri�Tethysand corresponding to the PETM interval, is not pro�nounced visually in the Nasypnoe section.

Methods

Three�step chemical treatment of palynologicalsamples using standard procedure of the Laboratory ofPaleofloristics, Geological Institute, Russian Acad�emy of Sciences (GIN), was applied. Each sample wasfirst treated in 10% HCl solution to dissolve carbonatecomponents of the rock, and then in a 5%Na2HPO4OH solution with subsequent removal ofclay minerals by decanting. Palynomorphs were sepa�rated from the precipitate by centrifuging in heavy liq�uid with a specific weight of 2.25 g/cm3 (KJ + CdJsolution) and collected into test tubes filled with glyc�erol for subsequent preparation of smear slides andmicroscopic examination. Rock samples and studiedsmear slides are hosted at the Laboratory of Paleoflo�ristics, GIN RAS, Moscow. Representative assem�blages of microphytoplankton (over 130 species intotal of dinocysts, acritarchs, prasinophytes, sporesand pollen of terrestrial plants) are identified in allsamples. Samples 19 and 20 were missed in palyno�morph study. The counte of not less than 200 palyno�morph specimens was made in each smear slide stud�ied. Biostratigraphic units were defined based onquantitative and taxonomic changes in palynomorphassemblages. Paleoenvironments were reconstructedfollowing Powell et al. (1996) from diagrams showingabundance ratios of different palynomorph groups anddinocyst ecogroups.

426


ALEKSANDROVA, SHCHERBININA

Lithology Biotic events in microphytoplankton assemblages

Nannoplankton Dinocysts

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Discoaster araneusD. anartios

Apectodinium augustumAcme Apectodinium

Senegalinium obscurum,Gen sp. Indet l Helm.�Cl., Costa, 1989

Rhomboaster spp.Discoaster cf. mahmoudiiD. anartiosD. araneus

Polysphaeridium zoharyApectodinium augustum,Muratodinium fimbriatum

Fasciculithus schaubiiF. thomasii,F. richardii,Heliolithus spp.Scapholithus apertusPlacozygus sigmoides

Pontosphaera planaRhabdosphaera sola

Discoaster multiradiatusRhomboaster intermediaradiation Fasciculithus

Heliolithus riedelii

Acme Apectodinium spp.Wilsonidium pechoricum

Apectodinium parvum,A. quinquilatum,Diphyes colligerum,Adnatosphaeridium sp. 1

Apectodinium homomorphumDeflandrea oebisfeldensisAdnatosphaeridium vittatum

Alisocysta margarita

Feodosiya

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Fig. 1. Geographic locality of the Nasypnoe section, lithology of the Paleocene–Eocene transition, and most significant bioticevents in nannoplankton and dinocyst assemblages: (1) Lithothamnium limestone, (2) bioclastic limestone, (3) sandy to silty clay,(4) clay.

Plate I. Nannofossils from the Paleocene–Eocene deposits of the Nasypnoe section:(1) Chiasmolithus bidens, Sample 5; (2) C. bidens, the same sample, polarized light (PL); (3) C. bidens, Sample 1, PL; (4) Chias�molithus consuetus, Sample 6; (5) C. consuetus, the same sample, PL; (6) Chiasmolithus titus, Sample 5; (7) C. titus, the same sam�ple, PL; (8) Coccolithus pelagicus, Sample 6; (9) C. pelagicus, the same sample, PL; (10, 11) Discoaster anartios, Sample 10;(12) Coccolithus robustus, Sample 2; (13) C. robustus, the same sample, PL; (14) Campylosphaera eodela, Sample 6; (15) C. eodela,the same sample, PL; (16) Discoaster araneus, Sample 9; (17) Discoaster cf. mahmoudii, Sample 10; (18) Discoaster falcatus, Sam�ple 5; (19) Discoaster splendidus, Sample 5; (20) Discoaster lenticularis, proximal side, Sample 13; (21) D. lenticularis, distal side,Sample 14; (22) Discoaster mohleri, proximal side, Sample 1; (23) Discoaster mediosus, Sample 5; (24) Discoaster salisburgensis,lateral side, Sample 14; (25) D. salisburgensis, Sample 5; (26) Discoaster mohleri, distal side, Sample 15; (27) Discoaster multira�diatus, distal side, Sample 9; (28) D. multiradiatus, proximal side, Sample 7; (29) Ellipsolithus distichus, Sample 6; (30) E. disti�chus, the same sample, PL; (31) Ellipsolithus macellus, Sample 13, PL; (32) Rhabdosphaera sola, Sample 5; (33) R, sola, the samesample, PL; (34) Hornibrookina arca, Sample 6; (35) H. arca, the same sample, PL; (36) Scapholithus apertus, PL, Sample 5.



Plate I

5 µ

m

1 2 3 4 5

6 7

12 13

8 9

10 11

14 15 16 17

18

2122

19 20

23 2425

26

29

2827

31 32 33

30 34 35 36

428



Plate II. Nannofossils from the Paleocene–Eocene deposits of the Nasypnoe section:(1) Bomolithus conicus, Sample 1, PL; (2) Heliolithus kleinpellii, Sample 6; (3) H. kleinpellii, Sample 2, PL; (4) Heliolithus riedelii,Sample 2; (5) H. riedelii, the same sample, PL; (6) H. riedelii, lateral side, Sample 2; (7) H. riedelii, the same sample, PL;(8) Heliolithus universus, Sample 2; (9) Heliolithus universus, the same sample, PL; (10) Heliolithus universus, lateral side, Sample 1,PL; (11) Heliolithus sp., Sample 1, PL; (12) Heliolithus bukryi, lateral side, Sample 2; (13) H. bukryi, the same sample, PL;(14) Heliolithus sp., Sample 1, PL; (15) Heliolithus sp., Sample 7, lateral side; (16) Heliolithus sp., the same sample, PL; (17) Fas�ciculithus tonii, Sample 6; (18) F. tonii, the same sample, PL; (19) Fasciculithus mitreus, Sample 6, PL; (20) Fasciculithus involu�tus, Sample 2, PL; (21) F. involutus, the same sample, PL; (22) Fasciculithus cf. involutus, Sample 2, PL; (23) Fasciculithusschaubii, Sample 3, PL; (24) Fasciculithus thomasii, Sample 3; (25) F. thomasii, the same sample, PL; (26) Fasciculithus aubertae,Sample 1, PL; (27) Fasciculithus sidereus, Sample 3, PL; (28) Fasciculithus tympaniformis, Sample 10, PL; (29) Fasciculithusbobii, Sample 1, PL; (30) Fasciculithus sp. 1, Sample 3, PL; (31) Fasciculithus sp. 2, Sample 3; (32) Fasciculithus sp. 2, the samesample, PL; (33) Fasciculithus sp. 3, Sample 2, PL; (34) Sphenolithus primus, Sample 6, PL; (35) Toweius sp., Sample 14, PL; (36)Toweius pertusus, Sample 15; (37) T. pertusus, the same sample, PL; (38) Toweius eminens, Sample 1; (39) T. eminens, the samesample, PL; (40) Toweius tovae, Sample 7; (41) T. tovae, the same sample, PL; (42) Toweius sp., Sample 3; (43) Toweius sp., thesame sample, PL; (44) Toweius occultatus, Sample 16, PL; (45) Toweius serotinus, Sample 5; (46) T. serotinus, the same sample,PL; (47) Neochiastozygus chiastus, Sample 2, PL; (48) N. chiastus, the same sample, PL; (49) Neochiastozygus concinnus, Sample 2;(50) N. concinnus, the same sample, PL; (51) Neochiastozygus denticulatus, Sample 15; (52) Neochiastozygus distentus, Sample 6;(53) N. distentus, the same sample, PL; (54) Neochiastozygus cf. distentus, Sample 6, PL; (55) Neochiastozygus junctus, Sample 9;(56) N. junctus, the same sample, PL; (57) Zygodiscus herlyni, Sample 5; (58) Pontosphaera plana, Sample 5; (59) Transversopontispulcher, Sample 10.

Nannofossils were studied in smear slides made bystandard technics (Bown, 1998) from all samples usingOlympus BX�41 microscope and photographing wasmade with Infinity X videocamera. The ratios of dif�ferent paleoecological groups in nannofossil assem�blages were statistically assessed by counting their spe�cies among 300 specimens in random fields of view.

Characteristic nannofossil and dinocyst taxa arefigured in Plates I–VI.

RESULTS AND DISCUSSION

Nannofossils

Despite mainly low carbonate content in the sam�pled clayey rocks, nannofossil assemblages are ratherdiverse (over 55 species in total have been identifiedwithin the sampled section interval) and characterizedby moderate to good preservation. Rare redepositedCretaceous nannofossil taxa are presented throughoutthe interval studued. In the Lithothamnium�bearinglimestone nannofossils were not found.

The nannofossil assemblage from basal part of thesandy�clayey sequence (samples 1 and 2, Plate 1) isrelatively abundant and diverse including commonCoccolithus spp., Prinsius martinii, diverse helioliths(Heliolithus kleinpellii, H. riedelii, H. bukryi, a. o.),frequent Toweius spp., Discoaster mohleri, and someother forms (Plates I and II). This assemblages is fea�tured for NP8 nannofossil zone. Total abundances ofcold�water chiasmoliths and warm�water discoastersare low and approximately equal (Fig. 2), that morelikely indicates temperate conditions. The abundanceof eurytopic Coccolithus spp. is higher than that ofeutrophic Toweius spp.

In Sample 3, total nannofossil abundance dropsand the ratio of ecologic groups changes significantly.The NP9 zonal marker Discoaster multiradiatus andRhomboaster intermedia appear at this level andremarkable fasciculith radiation with occurrence of

large daedal nannoliths (Fasciculithus schaubii, F. ala�nii, F. richardii, F. tonii, a.o.; Plate II), although theirtotal abundance remains to be almost the same. Theabundance Prinsius, Heliolithus, and, to a lesserextent, Coccolithus reduces, whereas the content ofDiscoaster spp. and Toweius spp. increases. In theSample 4, nannofossil abundance and diversity dra�matically decline, sediment becomes more silty andcontains glauconite that more likely evidences short�lived sea�level fall. Discoaster multiradiatus andToweius pertusus dominate low�abundant nannofossilassemblage. Upsection (samples 5–7), nannofossilsbecome to some extent recovered in abundance,although their total abundance remains to be relativelylow, but species diversity significantly increases. Newtaxa Rhabdosphaera sola and Pontosphaera plana firstappear and Zygodiscus herlyni, Neochiastozygus junc�tus, and Scapholithus apertus also first occur at thislevel. Such an early appearance if pontosphaerids andrhabdosphaerids in the middle of NP9 zone in its cur�rent understanding has never been reported from thenortheastern Peri�Tethys. Their appearance wasrecently considered to be related to the sapropeliticbed corresponding to PETM (N. Muzylev, pers. com�munication; Gavrilov et al., 2003). However, the firstPontosphaera spp. appear just in NP9 zone in the shelfsediments of New Jersey (Bybel and Self�Trail, 1995).A gradual increase in the nannofossil abundance anddiversity and appearance of new taxa are more likelyassociated with a rapidly progressing transgression andformation of new ecological niches.

The first occurrence of Rhomboaster spp.(Plate III), the marker of the PETM onset and lowerboundary of NP10, is found in Sample 8, where weidentified also the asymmetrical “excursion species”Discoaster anartios, D. araneus, and D. cf. mahmoudii(Plate I). Transversopontis pulcher also first appear atthis level, while Scapholithus apertus and Placozygussigmoides, which survived during the Creta�ceous/Paleogene extinction event, helioliths and all



Plate II

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6 7 8 9 10 11

12 13 14 15 16

17 18 19 20 21

23 24 25 26 27 28 29

30 31 32 33 34 3536

37

44

38 39 40 41 42 43 51

45 46 47 48 49 5057

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5952 53 54 5655

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22

430



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Plate III

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87 9 106

1312 14 1511

16 17 18 19 20

21 22 23 24 25

Plate III. Nannofossils from the Lower Eocene deposits of the Nasypnoe section:(1) Rhomboaster intermedia, Sample 7, PL; (2) R. intermedia, the same sample, PL; (3) R. intermedia, Sample 7; (4) R. intermedia,the same sample, PL; (5) Rhomboaster cf. cuspis, Sample 7, PL; (6) R. cuspis, Sample 10; (7) R. cuspis, the same sample, PL;(8) R. cuspis, Sample 10; (9) Rhomboaster bramlettei, Sample 13; (10) R. bramlettei, Sample 15; (11) R. cuspis, Sample 14;(12) R. cuspis, the same sample, PL; (13) R. calcitrapa, Sample 11; (14) Rhomboaster calcitrapa, Sample 9; (15) R. calcitrapa, thesame sample, PL; (16) Rhomboaster bitrifida, Sample 13; (17) R. bitrifida, the same sample, PL; (18) Rhomboaster spineus, Sam�ple 13; (19) Rhomboaster sp. 1, Sample 11; (20) Octolithus multiplus, Sample 12; (21) Rhomboaster bitrifida, Sample 13;(22) R. bitrifida, Sample 18; (23) Rhomboaster sp. 2, Sample 13; (24) nannolith, Sample 13; (25) the same sample, PL.

Plate IV. Dinocysts from the Paleocene deposits of the Nasypnoe section (magnification ×500 in all the figures):(1–3) Alisocysta margarita, Sample 1; (4) Florentinia ferox, Sample 2; (5) Cordosphaeridium gracile, Sample 1; (6) Areoligera gip�pigensis, Sample 1; (7) Cerodinium speciosum, Sample 1; (8) Spinidinium densispinatum, Sample 2; (9) Thalassiphora pelagica,Sample 2; (10) Melitasphaeridium pseudorecurvatum, Sample 2; (11) Cordosphaeridium fibrospinosum, Sample 1.



20 µm

Plate IV

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Plate V

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Plate V. Dinocysts from the Paleocene deposits of the Nasypnoe section (magnification ×500 in all the figures):(1–4) Phthanoperidinium crenulatum, Sample 3; (5) Adnatosphaeridium robustum, Sample 3; (6) Noremia sp., Sample 3; (7) Mel�itasphaeridium pseudorecurvatum, Sample 3; (8) Operculodinium sp,, Sample 3; (9, 14) Apectodinium homomorphum: (9) Sample 3,(14) Sample 4; (10) Cleistosphaeridium sp., Sample 5; (11) Polysphaeridium zoharyi, Sample 5; (12) Cordosphaeridium inodes,Sample 3; (13, 15) Kallosphaeridium spp., Sample 4; (16) Deflandrea oebisfeldensis, Sample 5; (17) Fibrocysta axiale, Sample 5;(18, 19) Adnatosphaeridium vittatum, Sample 5; (20) Operculodinium uncinispinosum, Sample 4.

larger fasciculiths which appeared at the base of theunderlying nannofossil zone, become disappeared atthe base of this interval. A bloom of warm�water dis�coasters in this interval is apparently an indication ofconsiderable warming (Fig. 3). As distinct from theother sections of the northeastern Peri�Tethys(Gavrilov et al., 2003), where high abundance of dis�coasterids is recorded throughout the PETM interval,their peak abundance recorded in the Nasypnoe sec�tion appeared to be very short�lived. Upsection, abun�dance of discoasters becomes lower and they are rep�resented mostly by “excursion species”. Contrarily,the abundance of Rhomboaster spp. increases upsec�tion, reaching maximal value at the level of Sample 13(Figs. 2 and 3), where this taxon and Toweius spp. rep�resent more than 80% of the nannofossil assemblage.Evidently, this level of distinct domination of rhom�boasters, where even eutrophic Toweius are relativelyreduced in abundance, corresponds to the PETM cul�mination, when environmental conditions were mostbaneful for nannoflora, although all negativelyaffected factors are still poorly understood. The lowabundance of calcareous dinocysts Thoracosphaeraspp. (Fig. 2) is also noteworthy, as this group is indica�tive of stress environments and frequently attains highabundance in habitats unfavorable for nannoflora.Angori et al. (2007) showed that their content is rela�tively high in the PETM interval of pelagic sedimen�tary successions. This specific nannofossil assemblageremains up to the level of Sample 17 and species diver�sity and ratio of taxa reach pre�PETM values in theuppermost part of the section studied (Sample 18).Total nannofossil abundance persists to be relativelylow at this level, that is more likely related to thebeginning of sea�level fall. Species composition ofnannofossil assemblage, however, differs from the pre�crisis nannoflora. Besides characteristic Paleocenetaxa that became extinct at the PETM level, fascicu�liths and Coccolithus robustus disappear also. In thehemipelagic settings, fasciculiths become graduallyreplaced by oligotrophic species Zygrhablithus bijuga�tus (Giusberti et al., 2007; Monechi and Angori, 2006;Tremolada and Bralower, 2004), which is mostlyabsent at the Paleocene–Eocene transition of thenortheastern Peri�Tethys. It has not been found in theNasypnoe section either. In the uppermost part ofinterval studied, rhomboasters decrease in abundance,but both short�arm and long�arm forms persist to bepresented. The pre�crisis Coccolithus/Toweius ratio isrestored here, and small T. pertusus/callosus signifi�cantly dominate among Toweius. The re�entry of nan�nofossil taxa that populated the basin before the crisis

most likely indicates the post�PETM restoration ofmesotrophic conditions.

Based on the compositional variations in the nan�nofossil assemblages, we recognize successive stages ofpaleoecological changes recorded in the Paleocene–Eocene transitional beds in the section studied. Thefirst stage showing the appearance of the largest dis�coasters (D. multiradiatus), fasciculiths and first rhom�boasters corresponds to beginning of NP9 zone. Firstcrisis in the nannofossil distribution featured byreduced abundance and diversity of nannofossils isfound in the middle part of NP9. Later (upper part ofNP9 zone), new nannofloral bloom occurs showingthe appearance of the new genera Pontosphaera andRhabdosphaera. This wellbeing was interrupted by achange in composition of nannoplankton communi�ties, when many earlier existed taxa became extinctand several short�lived species appear. The range ofnannofossil assemblage corresponding to the PETMcovers 4 m of the section. Initial stage of this event ismarked by a widespread occurrence of warm�waterdiscoasters, whereas its culmination corresponds tothe level of maximal abundance of rhomboasters. Thesubsequent gradual decline of rhomboasters was asso�ciated with progressing restoration of former ratiosbetween different taxa. First indication of such a resto�ration is recorded already in upper part of the range of“excursion assemblage”. Thus, the PETM onset ismarked in the section by dramatic compositionalchanges in nannofossil assemblages, whereas recoveryphase appears to be very slow and gradual.

Dinocysts

The exposed lower part of the sandy�clayeysequence (samples 1 and 2) corresponds to the earlyThanetian NP8 nannofossil zone (Fig. 1, Plate 2) andis characterized by occurrence of important strati�graphicaly dinocyst taxa, such as Cerodinium specio�sum, Alisocysta margarita, and Phthanoperidimumcrenulatum that enable the correlation of this intervalto the Alisocysta margarita Zone of the West Europeandinocyst zonation (Powell, 1992; Powell et al., 1996)and Zone DP2b recognized in the Crimea (Andreeva�Grigorovich and Shevchenko, 2007). The composi�tion of dinocyst assemblage is similar to that from thePegwell Marls in the lower part of the Thanet SandFormation at the 26N Chron (Powell et al., 1996).

In Sample 3, first occurrence (FO) of Apectodiniumhomomorphum is defined. This species marks lowerboundary of the Apectodinium hyperacanthum Zone

434



Plate VI

20 µm

1 23

45

6

8 9 10

11 12

1314

15

16

7



Plate VI. Dinocysts from the Paleocene–lower Eocene deposits of the Nasypnoe section (magnification ×500 in all the figures):(1–3) Adnatosphaeridium sp. 1: (1) Sample 6; (2, 3) Sample 7; (4) Lanternosphaeridium lanosum, Sample 7; (5) aff. Kenleya sp.,Sample 8; (6, 12) Wilsonidium pechoricum: (6) Sample 7, (12) Sample 16; (7) Apectodinium quinquelatum, Sample 7; (8) Batia�casphaera sp, Sample 10; (9) Schizosporis reticulatus, Sample 7; (10) Reticulatosphaera actinocoronata, Sample 10; (11, 15, 16)Glaphyrocysta–Areoligera�group, Sample 8; (13) Apectodinium augustum, Sample 9; (14) Gen sp. indet 1 sensu Heilmann�Clausen et Costa, 1989, Sample 17.

in West European sections (Powell, 1992; Powell et al.,1996), DP3 zone in Crimea (Andreeva�Grigorovichand Shevchenko, 2007), standard D4c zone (Luter�bacher et al., 2004), Viborg 5 zone in Denmark (Heil�mann�Clausen, 1994), and DP6a subzone of theNorth Sea (Mudge and Bujak, 1996). In the Nasypnoesection, this zone correlates to the late Thanetian NP9zone and span interval from Sample 3 to Sample 7.Besides the FO of zonal marker, FOs ofAdnatosphaeridium vittatum, A. sp. 1., Diphyes col�ligerum, Hystrichokolpoma sp., and Operculodiniumseverinii are recorded in this interval. Dinocyst assem�blage of lower part of this interval is characterized bysignificant amount of Spiniferites spp. and Achomo�sphaera spp., but upward the give the way to differentspecies of Apectodinium genus and Deflandrea oebi�sfeldensis. Diversity and abundance of Apectodiniumincrease upsection reaching 20% at the top of thiszone. The FO of Wilsonidium pechoricum is found inSample 7. Single redeposited Cretaceous spores andpollen and rare dinocysts are observed. The maximumabundance of redeposited palynomorphs (10%) isdetected at the base of A. hyperacanthum zone.

The higher interval of the section (samples 8�17)corresponds to the Apectodinium augustum zone cor�related to the early Eocene NP10 nannofossil zone.The dinocyst zone ranges from the FO of Apectodin�ium augustum (Sample 9) to the last occurrence (LO)of this species (Sample 17) and corresponds to theApectodinium acme. Despite the absence of markerspecies in the Sample 8, abundance of Apectodiniumexceeds 30% in the dinocyst assemblage that meansthe onset of the Apectodinium acme and can be consid�ered as a criterion defining lower boundary of the rel�evant zone (Heilmann�Clausen, 1985). In this sec�tion, Apectodinium augustum range cover intervalbetween Samples 9 and 17. Recent study in differentworld regions showed that FO of A. augustum andApectodinium acme are associated with the negativecarbon isotope excursion (CIE) related to the PETMand defining the Paleocene–Eocene boundary(Crouch et al., 2001, 2003; Luterbacher et al., 2004;Aubry et al., 2007).

Considerable peak of abundance of Areoligera spp.(49%) and Glaphyrocysta spp. is detected in the Sam�ple 8 and another abundance peak of these taxa (33%)is found in the Sample 12. Within Apectodiniumaugustum zone, FOs of Fibrocysta essentialis,Polysphaeridium cf. zoharyi, Adnatosphaeridium multi�spinosum, A. reticulense, A. robustum, and Kenleya�group (Kenleya cf. pachycerata, aff. Kenleya sp.,

Muratodinium fimbriatum, Lanternosphaeridium lano�sum) are recorded. Appearance of these taxa in thesame zone is typical in sections of different regions ofAsia and Europe (Iakovleva et al., 2001; Crouch et al.,2003; Iakovleva and Heilmann�Clausen, 2007).

Diversity of dinocysts decreases in Sample 18,abundance of Apectodinium spp. consists 23% anddiverse Spiniferites species become to dominate thatmakes this assemblage similar to late Thanetian one. Itis noteworthy that dinocyst assemblages recognizedabove the Paleocene–Eocene transition in differentregions varies in composition and stratigraphic rangethat more likely reflects specific hydrological regimesof the basins. In most basins, a regressive episode (ero�sion and/or change in lithology) is detected. Forinstance, Zone E1 established in the North Sea sedi�ments above the A. augustum Zone corresponds to theD. oebisfeldensis/Glaphyrocysta ordinata peak andcharacterizes the so�called North Sea biotic crisis(Mudge and Bujak, 1996). Similar distribution patternof dinocysts has been observed in sections of the Lon�don basin (Powell et al., 1996), Volga regions (Ale�ksandrova and Radionova, 2006), and the Trans�Urals(Vasil’eva, 2000). As dinocys assemblage of Nasypnoesection is not similar to any known from other areas,we can provisionally only correlate this interval to theHystrichosphaeridium tubiferum Zone (E1) of theNorth Sea (Bujak and Mudge, 1994).

The ratio of different palynomorph groups variesthroughout the section (Figs. 4 and 5). The palyno�spectra of the lowermost Alisocysta margarita zone(samples 1 and 2) are dominated by dinocysts (about80%) including the diverse Spiniferites, representing56% of all dinocysts, and other goniaulacoid taxa thatis characteristic of neritic open marine habitats. Sig�nificant amount of Incertae sedis sp. 1 sensu Heil�mann�Clausen, 1985 (9%) is found in the Sample 1that most likely suggests fresh water input into surfacewater of the basin.

The upper Thanetian deposits of the Apectodiniumhyperacanthum Zone (samples 3�7), shows a gradualincrease in abundance of spores and pollen of higherplants with their maximum concentration at theuppermost part of this interval (43%; Sample 7).Abundance of heterotrophic taxa (Apectodinium spp.,Deflandrea�group) and green algae accepted as indi�cators of lowered salinity and nutrient�rich environ�ment (Sluijs et al., 2005) increases simultaneously.Goniaulacoid dinoflagellate cysts show a generaldecline accompanied by an increase in the abundanceof Operculodinium spp. and Cordosphaeridium�group

436



indicating relatively shallower habitats. Progressinggrowth in abundance of subtropical Apectodiniumgenus characterizes the late Thanetian warming(Brinkhuis et al., 1994). Among spores and pollen ofhigher plants, spores and angiosperm pollen quantita�tively predominate with gymnosperm pollen being lessfrequent. Abundance ratios of this kind among the ter�restrial palynomorphs suggest a lowland character oflandscapes in surrounding land areas. Hence, thedynamics of changes in palynomorph assemblage fromthe base upward in the section evidence a successiveshallowing of the basin during accumulation of the lateThanetian sediments and lowering of the base level oferosion in adjacent land.

In the Apectodinium augustum Zone (Sample 8)corresponding to the PETM onset, important bioticturnover is recorded, and dinocysts again representmore than 80% of all palynomorphs. In greater part ofthis zone (samples 8�15), dinocyst ecogroups are ofvariable composition and characterize unstable hydro�logical conditions during sedimentation. Prevailingheterotrophic taxa belong to the genera Apectodiniumand Deflandrea, Wilsonidium pechoricum, and new,constantly occurring Kenleya�ecogroups. The Apecto�dinium peak has been regarded as interrelated not onlywith higher temperature, but also with considerablyhigher concentration of nutrients in seawater (Powellet al., 1996; Crouch et al., 2001, 2003; Brinkhuis et al.,2006). It is acknowledged as well that dinocysts of thisgenus are tolerant to significant salinity fluctuations,because their peak is recorded in sediments depositedin basins with lowered salinity and in open oceans. Thehigh specific diversity of the genus and the Apectodin�ium peak indicate a highstand (Sluijs et al., 2007).

In deposits of the Apectodinium augustum Zone,the relative abundances of Spiniferites and Cor�dosphaeridium neritic groups are comparatively lowsuggesting that they accumulated in proximity to thecoastline during the PETM.

In the PETM epoch, the most distinct changes arerecorded in the Deflandrea�group. The Areoligera�ecogroup of presumably photosynthetic dinocystsdominating in basal part of Apectodinium augustumzone (Sample 8) is suggested to be indicator of high�energy coastal habitats (Powell et al., 1996) and evi�dences the transgressive pulse the PETM onset. Acomparable situation has been described from NewZealand, where Crouch et al. (2003) studied dinocystsfrom the CIE interval of the Tawanui section andinterpreted the high relative abundance of Glaphyro�cysta as a result of taxa transportation from the innerneritic zone of the basin. However, highly abundantAreoligera�group (Fig. 5) oppressed mainly het�erotrophic dinocysts (Apectodinium spp., Deflandrea�group, Wilsonidium pechoricum) that is difficult toexplain exceptionally by taxa transportation from ner�

itic zone of the basin. Another peak of the Areoligera�group abundance (33%) and simultaneous high rela�tive abundance (13%) of green algae and acanthomor�phic acritarchs are recorded in Sample 12 that enableto suggest two transgression phases across the PETMtransition in the Eastern Crimea.

A high relative abundance of Operculodinium�group (18.5%) in the Sample 15 which indicates shal�lower environment, can suggest reative shoaling.

Spores prevailing among palynomorphs of higherplants and low content of angiosperm and gymno�sperm pollen suggesting widespread swamp areas onthe surrounding land at the time of the Paleocene–Eocene transition.

Palynomorph assemblages from uppermost sam�ples 16 to 18 suggest accumulation in normal marineconditions of a shelf platform.

CONCLUSIONS

Our study of nannofossils and dinocysts from thePaleocene–Eocene transition of Nasypnoe sectionrevealed the occurrence of sediments accumulatedduring the global biotic crisis associated with PETM.The distribution of the “excursion taxa” of bothmicroplanktonic groups and fluctuations in their rela�tive abundance in the interval from Sample 7 to Sam�ple 17 and suggest that this part of section correspondsto the PETM. Despite some difference between envi�ronmental trends inferred from analysis of nannofossiland dinocyst variations, several successive stages in thebasin development can be recognized (Fig. 6).

Sediments of lower part of the sandy�clayeysequence (NP8 and Alisocysta margarita zones) wereaccumulated in neritic settings of a relatively coolmesotrophic basin populated by relatively abundantnannoflora and dinoflagellates communities.

In basal sediments of the NP9 and Apectodiniumhyperacanthum zones, significant changes in compo�sition of both microphytoplankton groups are defined.Within the upper Thanetian interval, abundance anddiversity of dinocysts representing the warm�waterheterotrophic genus Apectodinium increase upsectionand suggest eutrophication and warming in the basinduring this epoch. While the changes in palynomorphassemblages of the pre�PETM interval show a stabletrend of gradual sea level fall with the regression max�imum near the upper zonal boundaries, the nannofos�sil variations seems to be more complex.

Below the PETM interval there are two levels ofincreased abundance and diversification of nannofos�sils with intervening episode of a sharp decline inabundance and diversity (Sample 4) that most likelyindicates drastic sea level fall. In the dinocyst assem�blages, extremely low content of Deflandrea�grouponly corresponds to this nannofossil event. This sce�nario of dynamics in nannoplankton evolution differs



7 6 5 4 3

10 9 8 7 6 5 4 3 2 1 0

18 17 16 15 14 13 12 11 10 9 8

010

2030

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60A

vera

ge p

op

ula

tio

n o

f n

ann

op

lan

kto

n

Am

ou

nt

of

spec

ies

Chias

mol

ithus

spp.

Disc

oaste

r spp.

Cocco

lithu

s spp.

Towei

us sp

p.

Fascic

ulith

us sp

p.

Sphen

olith

us sp

p.

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cosp

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a sp

p.

Rhom

boas

ter s

pp.

00

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1010

2020

3030

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7 6 5 4 3 2 1

010

200

1020

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1020

3040

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ThanetianYpresian

NP8NP9NP10

Fig

. 2. V

aria

tion

s in

rel

ativ

e ab

unda

nce

s of

nan

nop

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ical

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nt o

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cies

(da

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r av

erag

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e) c

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in 5

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icro

scop

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w 0

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m in

dia

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er. T

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as

pale

oeco

logi

cal m

arke

rs a

re c

ool�

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er C

hias

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ithu

s spp

.; w

arm

�wat

er D

isco

aste

rsp

p.; e

uryt

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cos

mop

olit

an C

occo

lithu

s spp

.; e

utro

phic

Tow

eius

spp

. an

d F

asci

culit

hus s

pp.;

war

m�w

ater

oli

gotr

oph

ic S

phen

olit

hus s

pp.;

Tho

raco

spha

era

spp.

indi

cati

ve o

f un

fa�

vora

ble

con

diti

ons;

an

d sh

ort�

live

d R

hom

boas

ter

spp.

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cifi

c of

th

e P

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bols

for

lith

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y as

in F

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438



18

17

16

15

14

1312111098

0 10

765

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3

2

1

NP

8N

P9

NP

10

20

0 10 20 30 0 10 20 30 40 50 60 70 80 90

Discoaster spp.(“non�excursion”),

N/mm2

Discoaster spp.(“excursion”),

N/mm2Rhomboaster spp., N/mm2

from the eustatic trend established in the easternregions of the northeastern Peri�Tethys (Caucasus andCentral Asia), where sea level fall immediately pre�dates the deposition of the corresponding to PETMsapropelitic horizon. In the Nasypnoe section, theregression episode and biotic crisis are separated by aconsiderable time span, when changes in compositionof nannofossil assemblages were associated withappearance of new genera Pontosphaera and Rhab�dosphaera which correspond to the PETM interval inother sections. This can be explained by the muchgreater thickness of the PETM deposits in the Nasyp�noe section accumulated under conditions ofenhanced influx of siliciclastics. On the other hand,this discrepancy might be explained by the presence oftwo short�term regressive pulses, which could be rec�ognized in this shallow�water section only.

Sediments of the PETM interval accumulated dur�ing distinct transgression with two episodes of thebasin deepening that is consistent with the scenario ofthe accumulation of organic�rich deposits in the

northeastern Peri�Tethys (Gavrilov and Shcherbinina,2004). In the Nasypnoe section, deposits of this inter�val contains all taxa, which are characteristic of nan�nofossil and dinocysts assemblages from the PETMrecord in the Peri�Tethys and other regions of theWorld. These are the short�lived “excursion” speciesof nannofossils Rhomboaster spp., asymmetric Dis�coaster anartios, D. araneus, and dinocysts of the sub�tropical genus Apectodinium, including its speciesA. augustum which is typical for this event. The warm�water dinocysts dominating the assemblage across thePETM interval imply a considerable warming in thebasin, but a low relative abundance of discoastersremains poorly understood. A high abundance ofeutrophic and stress species most likely suggest anexcessive influx of nutrients into the basin, which pro�duced negative impact on most nannoplankton anddinoflagellate taxa.

The restoration of biota after the crisis was gradualand nannofossils remain to be of low abundance thatcan indicate shallow depth of the basin after PETM

Fig. 3. Variations in abundance (amount of specimens per 1 mm2 of preparation) of “non�excursion” Discoaster multiradiatus,D. falcatus, D. splendidus, D. mediosus, and “excursion” Discoaster anartios, D. araneus and Rhomboaster spp. Symbols for lithol�ogy as in Fig. 1.



Tab

le 1

.D

istr

ibut

ion

an

d ab

unda

nce

rat

es o

f pri

nci

pal n

ann

opla

nkt

on s

peci

es in

th

e N

asyp

noe

sec

tion

Nannoplankton zones

Sample nos.Chiasmolithus bidensC. eograndisCoccolithus pelagicusC. robustusDiscoaster mohleriFasciculithus aubertaeF. involutusF. tympaniformisHeliolithus kleinpelliiH. riedeliiNeochiastozygus concinnusPrinsius martiniiSphenolithus primusToweius eminensT. occultatusT. pertususDiscoaster falcatusEllipsolithus distichusLophodolithus nacensN. chiastiusToweius occultatusD. multiradiatusDiscoaster splendidusFasciculithus alaniiF. richardiiF. sidereusF. schaubiiF. thomasiiToweius tovaeChiasmolithus consuetusDiscoaster lenticularisD. salisburgensisEllipsolithus macellusNeochiastozygus distentusPontosphaera planaRhabdosphaera solaScapholithus apertusZygodiscus herlyniiChiasmolithus titusToweius serotinusNeochiastozygus junctusHornibrookina arcaNeochiastozygus concinnusCampylosphaera eodelaRhomboaster intermediaR. bramletteiD. anartiosD. araneusD. cf. mahmoudiiRhomboaster calcitrapaR. spineusR. cuspisTransversopontis pulcher

NP10

18f

cf

ff

ff

ff

ff

ff

ff

fr

ff

ff

r

17f

ff

ff

ff

ff

ff

ff

ff

ff

ff

16f

ff

ff

rf

ff

ff

ff

ff

ff

15f

ff

ff

cf

ff

ff

ff

fr

ff

14f

ff

ff

fc

ff

ff

ff

fr

f

13f

ff

ff

cf

ff

ff

fr

ff

cf

f

12f

ff

fc

ff

ff

ff

ff

ff

r

11r

ff

ff

cf

ff

ff

ff

ff

ff

cf

10f

ff

ff

cf

ff

ff

ff

cf

f

9f

ff

fr

cr

ff

fr

rf

rf

f

8f

ff

ff

fc

ff

rf

ff

f

NP9

7f

rf

ff

ff

ff

fc

ff

ff

ff

ff

ff

ff

ff

ff

ff

ff

6f

fr

ff

ff

ff

ff

rf

fr

ff

fr

ff

ff

ff

ff

ff

ff

ff

ff

ff

5f

ff

ff

rf

fc

ff

ff

ff

ff

ff

ff

ff

ff

ff

ff

f

4f

ff

ff

ff

ff

ff

f

3f

ff

rf

ff

ff

ff

ff

fr

ff

ff

ff

f

NP8

2f

cr

ff

ff

ff

ff

ff

ff

ff

f

1f

fc

ff

ff

ff

ff

rf

ff

ff

Not

e:(f

) fe

w (

1 to

10

spec

imen

s pe

r a

mou

nt; (

r) r

are

(1 s

peci

men

at l

east

in e

very

sec

ond

fiel

d of

vie

w);

(c)

cro

wde

d (u

p to

5 s

peci

men

s in

eve

ry fi

eld

of v

iew

).

440



Tab

le 2

.D

istr

ibut

ion

an

d am

oun

t of

pal

ynom

orph

s in

th

e N

asyp

noe

sec

tion

Sam

ple

nos.

Tax

a1

23

45

67

89

1011

1213

1415

1617

18

Ach

omos

phae

ra sp

p.32

1427

3323

3210

27

813

76

55

207

Spin

iferi

tes s

pp.

208

7111

474

5852

374

2829

3943

3548

3519

6177

Cor

dosp

haer

idiu

m d

iver

gens

e E

isen

ack,

196

37

312

44

25

31

Cor

dosp

haer

idiu

m in

odes

(K

lum

pp)

Eis

enac

k, 1

963

106

811

26

32

13

23

65

Cor

dosp

haer

idiu

m fi

bros

pino

sum

Dav

ey e

t Will

iam

s, 1

966

31

Cor

dosp

haer

idiu

m g

raci

le (

Eis

.) D

avey

et W

illia

ms,

196

64

13

12

11

Olig

osph

aeri

dium

com

plex

(W

hite

) D

avey

et W

illia

ms,

196

64

Con

nexi

mur

a fim

bria

ta (

Mor

genr

oth)

May

, 198

02

12

Haf

nias

phae

ra se

ptat

a (C

ook.

et E

is.)

Han

sen,

197

715

52

102

62

Gla

phyr

ocys

ta c

f. or

dina

ta (

Dav

. et W

ill.)

Sto

ver e

t Evi

tt, 1

978

44

Gla

phyr

ocys

ta sp

.1

Are

olig

era

seno

nens

is L

ejeu

ne�C

arpe

ntie

r, 19

384

1

Are

olig

era

gipp

igen

sis J

olle

y, 1

992

1

Are

olig

era

sp.

51

1

Alis

ocys

ta m

arga

rita

Har

land

, 197

95

71

2

Alis

ocys

ta sp

. 2 se

nsu

Hei

lman

n�C

laus

en, 1

985

31

1

Dip

hyes

spin

ulum

(D

rugg

) St

over

et E

vitt

, 197

815

3

Dip

hyes

sp.

2

Bat

iaca

spha

era

spha

eric

a St

over

, 197

77

3

Bat

iaca

spha

era

sp.

234

7

?Sur

culo

spha

erid

ium

sp.

3

Par

alec

anie

lla in

dent

ata

(Def

l. et

Coo

k.)

Coo

k. e

t Eis

., 19

702

12

11

Ope

rcul

odin

ium

unc

inis

pino

sum

(de

Con

inck

) Is

lam

, 198

39

216

167

512

817

911

Impa

gidi

nuim

sp. 1

sens

u H

eilm

ann�

Cla

usen

, 198

53

6

Com

etod

iniu

m?

com

atum

Sri

vast

ava,

198

411

48

11

1

Mel

itasp

haer

idiu

m p

seud

orec

urva

tum

(M

orge

nr.)

Buj

ak, 1

980

67

66

21

1

Cer

odin

ium

spec

iosu

m (

Alb

erti)

Len

t. et

Will

., 19

875

151

Ince

rtae

sedi

s sp.

1 se

nsu

Hei

lman

n�C

laus

en, 1

985

422

inde

term

ined

cho

rate

cys

ts8

89

127

2719

8

Pal

aeot

etra

dini

um m

inus

culu

m (

Alb

erti)

Sto

ver e

t Evi

tt, 1

978

12

11

Ely

troc

ysta

bre

vis S

tove

r et H

ande

rbol

, 199

43

212

21



Tab

le 2

.(C

ontd

.)

Sam

ple

nos.

Tax

a1

23

45

67

89

1011

1213

1415

1617

18

Cal

igod

iniu

m a

mic

ulum

Dru

gg, 1

970

1

Kal

losp

haer

idiu

m b

revi

barb

atum

de

Con

inck

, 196

91

21

56

16

14

11

11

13

4

Cyc

lone

phel

ium

sp.

1

Cri

brop

erid

iniu

m sp

.2

12

11

12

Hys

tric

hosp

haer

idiu

m tu

bife

rum

(E

hren

berg

) D

efla

ndre

, 193

73

35

11

32

12

3

Spin

idin

ium

den

sisp

inat

um S

tanl

ey, 1

965

1

Pht

hano

peri

dini

um c

renu

latu

m (

de C

onin

.) L

ent.

et W

ill.,

1977

12

64

71

12

11

12

Pht

hano

peri

dini

um p

aleo

ceni

cum

Luc

as�C

lark

, 200

61

Spin

idin

ium

ech

inoi

deum

(C

ook.

et E

is.)

Len

t. et

Will

., 19

763

45

Tha

lass

ipho

ra p

elag

ica

(Eis

.) E

isen

ack

et G

ocht

, 196

01

1

Tha

lass

ipho

ra d

elic

ata

Will

iam

s et D

owni

e, 1

966

312

1

?Cer

ebro

cyst

a sp

.3

54

1

Cla

dopy

xidi

um sa

eptu

m (

Mor

genr

.) S

tove

r et E

vitt

, 197

81

1

Are

olig

era

coro

nata

(W

etze

l) L

ejeu

ne�C

arpe

ntie

r, 19

381

Ope

rcul

odin

ium

cen

troc

arpu

m (

Def

l. et

Coo

k.)

Wal

l, 19

671

617

222

51

47

2110

10

Tri

gono

pyxi

dia

gine

lla (

Coo

k. e

t Eis

.) D

own.

et S

arje

ant,

196

51

1

Alis

ocys

ta sp

. 1

Ape

ctod

iniu

m h

omom

orph

um (

Def

l. et

Coo

k.)

Len

t. et

Will

., 19

77

38

3516

61

7618

2924

3249

2241

4323

Def

land

rea

oebi

sfel

dens

is A

lber

ti, 1

959

24

2141

61

193

109

520

218

1715

Def

land

rea

sp.

25

41

Flo

rent

inia

fero

x (D

efla

ndre

) D

uxbu

ry, 1

980

2

Cri

brop

erid

iniu

m w

etze

lii (

Lej

eune

�Car

pent

ier)

Hel

enes

, 198

41

1

Cor

dosp

haer

idiu

m sp

.1

14

17

11

Are

olig

era

cf. s

enon

ensi

s Lej

eune

�Car

pent

ier,

1938

32

71

Spin

idin

ium

sp.

1

Pol

ysph

aeri

dium

cf.

subt

ile D

avey

et W

illia

ms,

196

62

1

Dip

hyes

cf.

appe

ndic

ular

e C

ooks

on e

t Eis

enac

k, 1

970

1

Adn

atos

phae

ridi

um v

ittat

um W

illia

ms e

t Dow

nie,

196

63

132

23

13

Impa

gidi

nium

cf.

velo

rum

Buj

ak, 1

984

21

1

Pal

aeoc

ysto

dini

um g

olzo

wen

se A

lber

ti, 1

961

1

Sene

galin

ium

sp.

1

442



Tab

le 2

.(C

ontd

.)

Sam

ple

nos.

Tax

a1

23

45

67

89

1011

1213

1415

1617

18

Cer

odin

ium

die

belii

(A

lber

ti) L

entin

et W

illia

ms,

198

71

Cer

odin

ium

sp.

52

11

Ape

ctod

iniu

m p

arvu

m (

Alb

erti)

Len

tin e

t Will

iam

s, 1

977

11

429

1420

510

84

64

1

Ape

ctod

iniu

m q

uinq

uila

tum

(W

ill. e

t Dow

n.)

Cos

ta e

t Dow

nie,

197

91

95

62

33

49

815

1

Ape

ctod

iniu

m sp

.3

1912

1933

3429

3818

4135

2329

5620

Adn

atos

phae

ridi

um sp

. 11

13

13

74

27

41

Adn

atos

phae

ridi

um ro

bust

um (

Mor

genr

oth)

de

Con

inck

, 197

51

16

41

1

Cri

brop

erid

iniu

m c

f. te

nuita

bula

tum

(G

erla

ch)

Hel

enes

, 198

42

Hys

tric

hoko

lpom

a sp

.1

12

1

Ope

rcul

odin

ium

seve

rini

i (C

ook.

et C

ranw

ell)

Isl

am, 1

983

12

11

31

32

Dip

hyes

col

liger

um (

Def

l. et

Coo

k.)

Coo

kson

, 196

55

52

11

12

35

3

Gla

phyr

ocys

ta p

astie

lsii

(Def

l. et

Coo

k.)

Stov

er e

t Evi

tt, 1

978

11

Gla

phyr

ocys

ta c

f. vi

cina

(E

aton

) St

over

et E

vitt

, 197

81

Lin

gulo

dini

um m

acha

erop

horu

m (

Def

l. et

Coo

k.)

Wal

l, 19

671

44

12

Kal

losp

haer

idiu

m c

apul

atum

Sto

ver,

1977

3

Cor

dosp

haer

idiu

m e

xilim

urum

Dav

ey e

t Will

iam

s, 1

966

11

Ape

ctod

iniu

m h

yper

acan

thum

(C

ook.

et E

is.)

Len

t. et

Will

., 19

774

34

1111

1017

1317

1613

912

5

Cer

odin

ium

spec

iosu

m g

labr

um (

Goc

ht)

Len

t. et

Will

., 19

878

21

11

5

Ope

rcul

odin

ium

mic

rotr

ianu

m (

Klu

mpp

) Is

lam

, 198

3 7

515

103

42

44

Ope

rcul

odin

ium

sp.

211

68

25

Impa

gidi

nium

sp.

11

83

21

3

Len

tinia

wet

zelii

(M

orge

nrot

h) B

ujak

in B

ujak

et a

l., 1

980

21

81

32

3

Sene

galin

ium

obs

curu

m (

Dru

gg)

Stov

er e

t Evi

tt, 1

978

110

211

27

Cle

isto

spha

erid

ium

sp.

11

Kal

losp

haer

idiu

m s

p.3

23

2

Wils

onid

ium

pec

hori

cum

Iak

ovle

va e

t Hei

lm.�

Cla

usen

, 200

74

21

21

519

119

242

Ape

ctod

iniu

m p

anic

ulat

um (

Cos

ta e

t Dow

n.)

Len

t. et

Will

., 19

7710

75

3

Hys

tric

hoko

lpom

a cf

. uni

spin

um W

illia

ms e

t Dow

nie,

196

61

13

Cor

dosp

haer

idiu

m fi

bros

pino

sum

Dav

ey e

t Will

iam

s, 1

966

22

1

Rot

tnes

tia b

orus

sica

(E

isen

ack)

Coo

kson

et E

isen

ack,

196

11

13

1

Ret

icul

osph

aera

act

inoc

oron

ata

(Ben

ed.)

Buj

ak e

t Mat

suok

a, 1

986

12

84

63

4



Tab

le 2

.(C

ontd

.)

Sam

ple

nos.

Tax

a1

23

45

67

89

1011

1213

1415

1617

18

Haf

nias

phae

ra fl

uens

Han

sen,

197

75

4

Pol

ysph

aeri

dium

sp.

11

1

Hys

tric

hoko

lpom

a cf

. sal

acia

Eat

on, 1

976

11

41

33

24

2

Hys

tric

hosp

haer

opsi

s cf.

ovum

Def

land

re, 1

935

11

42

11

21

Mem

bran

osph

aera

sp.

1

Gla

phyr

ocys

ta –

Are

olig

era

grou

p31

6440

3967

8849

642

29

Fib

rocy

sta

vect

ensi

s (E

aton

) St

over

et E

vitt

, 197

83

2

aff.

Ken

leya

sp.

33

54

25

21

11

Lan

tern

osph

aeri

dium

lano

sum

Mor

genr

oth,

196

61

137

21

21

Ape

ctod

iniu

m a

ugus

tum

(H

arla

nd)

Len

t. et

Will

., 19

816

51

32

11

310

Ken

leya

cf.

pach

ycer

ata

Coo

kson

et E

isen

ack,

196

54

1

aff.

Wils

onid

ium

pec

hori

cum

Iak

ovle

va e

t Hei

lm.�

Cla

usen

, 200

72

Mur

atod

iniu

m fi

mbr

iatu

m (

Coo

kson

et E

isen

ack)

Dru

gg, 1

970

12

51

51

41

Adn

atos

phae

ridi

um m

ultis

pino

sum

Will

iam

s et D

owni

e, 1

966

1

Pal

aeop

erid

iniu

m sp

.2

Cri

brop

erid

iniu

m te

nuita

bula

tum

(G

erla

ch)

Hel

enes

, 198

43

32

Def

land

rea

dent

icul

ata

Alb

erti,

195

91

Mel

itasp

haer

idiu

m a

ster

ium

(E

aton

) B

ujak

et a

l., 1

980

1

Fib

rocy

sta

esse

ntia

lis (

de C

onin

ck)

Bri

nkhu

is e

t Zac

hari

asse

, 198

85

21

12

Impa

gidi

nium

cor

nutu

m M

atsu

aka

et B

ujak

, 198

81

Lei

osph

aeri

dia

sp.

1

aff.

Bat

iaca

spha

era

sp.

1720

12

Impl

etos

phae

ridi

um sp

.4

92

11

Pol

ysph

aeri

dium

cf.

zoha

ry (

Ros

sign

ol)

Buj

ak e

t al.,

198

04

46

28

1

Apt

eodi

nium

sp.

11

Cri

brop

erid

iniu

m sp

. A se

nsu

Cro

uch,

200

32

Haf

nias

phae

ra g

raci

osa

Han

sen,

197

71

Gen

. sp.

Ind

et. 1

sens

u H

eilm

ann�

Cla

usen

et C

osta

, 198

92

22

25

Are

olig

era

cf. c

oron

ata

(Wet

z.)

Lej

eune

�Car

pent

ier,

1938

1

Tha

lass

ipho

ra sp

inife

ra (

Coo

k. e

t Eis

.) S

tove

r et E

vitt

, 197

82

Hys

tric

hoko

lpom

a cf

. rig

audi

ae D

efla

ndre

et C

ooks

on, 1

955

1

444



Tab

le 2

.(C

ontd

.)

S

ampl

e no

s. T

axa

12

34

56

78

910

1112

1314

1516

1718

Adn

atos

phae

ridi

um re

ticul

ense

(P

astie

ls)

de C

onin

ck, 1

969

1S

um45

919

526

925

724

724

820

814

140

622

833

929

027

331

026

325

239

424

3

Pra

sin

oph

ytes

, ac

rita

rch

s, a

nd

oth

er p

alyn

omor

phs

Aca

ntam

orh

acri

tarc

hs6

512

2614

1210

6M

ycrh

ystr

idiu

m sp

.2

1H

orol

ogin

ella

incu

rvat

a C

ooks

on e

t Eis

enac

k, 1

962

11

21

Fro

mea

laev

igat

a (J

iabo

) L

ent.

et W

ill.,

1981

11

Fro

mea

frag

ilis (

Coo

k. e

t Eis

.) S

tove

r et E

vitt

, 197

82

1P

alam

bage

s sp.

21

11

Gre

en a

lgae

56

2133

2524

421

62

12Ta

sman

ites s

p.1

3Sp

irog

yra

sp.

214

11

31

Pte

rosp

erm

ella

sp.

41

1N

orem

ia sp

.6

3L

eios

phae

ridi

a sp

.1

Scol

ecod

onts

11

Mic

rofo

ram

inife

ral l

inin

gs4

Sum

106

3443

2627

714

120

4827

225

141

6

Spo

res

and

poll

en o

f hig

her

pla

nts

Spo

res

2610

2768

111

6158

533

2234

3736

2132

2344

33P

olle

n of

gym

nosp

erm

4630

6655

118

6788

25

71

1214

54

711

24P

olle

n of

ang

iosp

erm

153

949

8048

379

819

2110

186

74

67

Sum

8743

102

172

309

176

183

1646

4856

5968

3243

3461

64

Red

epos

ited

pal

ynom

orph

s

Spo

res

and

poll

en o

f hig

her

pla

nts

147

3614

515

197

103

2L

itosp

haer

idiu

m a

rund

um (

Eis

. et C

ook.

) D

avey

, 197

91

Cri

brop

erid

iniu

m m

uder

ogen

se (

Coo

k. e

t Eis

.) D

avey

, 196

9 1

11

1T

rith

yrod

iniu

m sp

.1

Leb

erid

ocys

ta sp

.1

Apt

ea sp

.1

Alte

rbid

iniu

m a

cutu

lum

(W

ilson

) L

ent.

et W

ill.,

1985

1O

ligos

phae

ridi

um a

ster

iger

um (

Goc

ht)

Dav

ey e

t Will

., 19

691

Gon

yaul

acac

ysta

sp.

1O

dont

ochi

tina

oper

cula

ta (

Wet

zel)

Def

l. et

Coo

k., 1

955

1S

um18

837

155

1622

07

103

01

20

00

0



10 9 8 7 6 5 4 3 2 1 0

18 17 16 15 14 13 12 11 10 9 8

010

7 6 5 4 3 2 1

ThanetianYpresian

2030

4050

6070

8090

010

2030

4050

05

1015

05

1015

05

1015

200

510

150

510

15%

Co

nce

ntr

atio

ns

of

pal

yno

mo

rph

gro

up

s (%

rel

ativ

e to

all

sp

ecim

ens)

Din

ocy

sts

Sp

ore

s/p

oll

enP

rasi

no

ph

ytes

/ac

rita

rch

sS

po

res

of

hig

her

pla

nts

Po

llen

of

gym

no

sper

ms

Po

llen

of

angi

osp

erm

sR

edep

osi

ted

pal

yno

mo

rph

s

PALEOCENEEOCENESeries

Stage

Formation, sequence

Thickness, m

Sandy�argillaceous rocks

Alisocystamargarita

A. hypera�canthumA. augustumH. tubiferumDinocyst zones

Fig

. 4. C

once

ntr

atio

ns

of p

alyn

omor

ph g

roup

s (%

) at

th

e sa

mpl

ing

leve

ls. S

ymbo

ls fo

r li

thol

ogy

as in

Fig

. 1.

446



10 9 8 7 6 5 4 3 2 1 0

18 17 16 15 14 13 12 11 10 9 8

010

7 6 5 4 3 2 1

ThanetianYpresian

2030

4050

600

1020

3040

500

510

1520

05

10%

PALEOCENEEOCENESeries

Stage

Formation, sequence

Thickness, m

Sandy�argillaceous rocks

Alisocystamargarita

A. hypera�canthumA. augustumH. tubiferumDinocyst zones

010

2030

4050

05

1015

200

510

05

100

1020

3040

Spinife

rites

�gro

up

Areoli

gera

�gro

up

Apecto

diniu

m sp

p.

Deflandre

a�gro

up

Operc

ulodin

ium

spp. Cor

dosph

aerid

ium

�gro

up

Kenley

a�gro

up Wils

onid

ium

pech

oricu

m

other

Gonya

ulacoid

Fig

. 5. C

once

ntr

atio

ns

of e

colo

gic

din

ocys

t gr

oups

(%

rel

ativ

e to

all

din

ocys

ts)

at t

he

sam

plin

g le

vels

. Sym

bols

for

lith

olog

y as

in F

ig. 1

.



10

9

8

7

6

5

4

3

2

1

0

18

17

16

15

14

1312111098765

4

3

2

1

NP

8N

P9

NP

10

PA

LE

OC

EN

EE

OC

EN

E

Sea�levelchanges

Basin conditions

Temperature Trophicity degree

Regression

Transgression

?

Regression

High sealevel

pre

�PE

TM

Onset

Peak

Res

tora

tio

n

PE

TM

Cool

Warm

Mesotrophic

Eutrophic

Supereutrophic?

Mesotrophic

Eutrophic

Warm

Cool

Cool

Fig. 6. Interpretation of the PETM paleoenvironments based on nannofossils and dinocysts. Symbols for lithology as in Fig. 1.

termination. It is supported also by relatively increasedcontent of spores and pollen and relative decrease inabundance of dinocysts in the uppermost part of thesection. At the same time, the restoration of dinocystwas accompanied by re�entry of all taxa, while nanno�fossil assemblages appeared to be substantiallychanged and lost many typically Paleocene taxa as aresult of PETM impact.

ACKNOWLEDGMENTS

The work was supported by the Russian Founda�tion for Basic Research, project no. 09�05�00872, andby the Program of Presidium RAS no. 24.

Reviewers E.M. Bugrova and M.A. Akhmetiev

REFERENCES

Akhmetiev, M.A. and Zaporozhets, N.I., Succession ofDinocysts in the Paleogene and Lower Miocene Sections ofthe Russian Platform, Crimea–Caucasus Region and TuranPlate as Evidence of Ecosystem Reorganizations, in Isko�paemye organizmy kak osnova stratigrafii, korrelyatsii i pale�obiogeografii fanerozoya (Fossil Organisms as Basis of Phan�erozoic Stratigraphy, Correlation and Paleobiogeography),Moscow: GEOS, 1996, pp. 55–69.Aleksandrova, G.N. and Radionova, E.P., On the LatePaleocene Stratigraphy of the Saratov Volga Region: Micro�

paleontological Characteristics of the Kamyshin Forma�tion, Dyupa Gully Section, Paleontol. J, 2006, vol. 40,suppl. 5, pp. 543–557.Andreeva�Grigorovich, A.S. and Shevchenko, T.V.,Dinocyst Zonation in the Paleocene Deposits of Ukraine,in Paleontol. doslidzhennya v Ukraini (Paleontological Stud�ies in Ukraine), Kiev: IGN NANU, 2007, pp. 211–214.Angori, E., Bernaola, G., and Monechi, S., CalcareousNannofossil Assemblages and Their Response to the Pale�ocene–Eocene Thermal Maximum Event in Different Lat�itudes: ODP Site 690 and Tethyan Sections, Spec. Pap. Geol.Soc. Am., 2007, no. 424, pp. 69–85.Aubry, M.P., Ouda, K., Dupuis, C., et al., The Global Stan�dard Stratotype Section and Point (GSSP) for the Base ofthe Eocene Series in the Dababaya Section (Egypt), Epi�sodes, 2007, vol. 3, no. 4, pp. 271–286.Bown, P.R., Calcareous Nannofossil Biostratigraphy, BritishMicropalaeontol. Soc. Publ. Ser., London: Chapman andHall Ltd., Kluwer Academic Publisher, 1998.Brinkhuis, H., Romein, A.J.T., Smit, J., and Zacariasse, J.�W.,Danian–Selandian Dinoflagellate Cysts from Lower Lati�tudes with Special Reference to the El Kef Section, NWTunisia, GFF, 1994, vol. 116, pp. 46–48.Brinkhuis, H., Schouten, S., Collinson, M.E., et al., Epi�sodic Fresh Surface Waters in the Eocene Arctic Ocean,Nature, 2006, vol. 441, pp. 606–609.Bugrova, E.M., Zakrevskaya, E.Yu., and Tabachnikova, I.P.,New Data on Paleogene Biostratigraphy of the EasternCrimea, Stratigr. Geol. Korrelyatsiya, 2002, vol. 10, no. 1,

448



pp. 83–93 [Stratigr. Geol. Correlation (Engl. Transl.),vol. 10, no. 1, pp. 77–87].

Bujak, J. and Mudge, D., A High�Resolution North SeaEocene Dinocyst Zonation, J. Geol. Soc., 1994, vol. 151,pp. 449–462.

Bybel, L.M. and Self�Trail, J.M., Evolutionary, Biostrati�graphic, and Taxonomic Study of Calcareous Nannofossilsfrom a Continuous Paleocene–Eocene Boundary Sectionin New Jersey, Prof. Pap. US Geol. Surv., 1995, no. 1554,pp. 1–112.

Clyde, W.C. and Gingerich, P.D., Mammalian CommunityResponse to the Latest Paleocene Thermal Maximum: AnIsotaphonomic Study in the Northern Bighorn Basin, Wyo�ming, Geology, 1998, vol. 26, pp. 1011–1014.

Crouch, E.M., Heilmann�Clausen, C., Brinkhuis, H.,et al., Global Dinoflagellate Event Associated with the LatePaleocene Thermal Maximum, Geology, 2001, vol. 29,pp. 315–318.

Crouch, E.M., Brinkhuis, H., Visscher, H., et al., LatePaleocene–Early Eocene Dinoflagellate Cyst Record fromthe Tethys: Further Observations on the Global Distribu�tion of Apectodinium, in Causes and Consequences of Glo�bally Warm Climates in the Early Paleogene, Wing S.L.,et al., Eds., Spec. Pap. Geol. Soc. Am., 2003, no. 369,pp. 113–131.

Gavrilov, Yu.O. and Shcherbinina, E.A., Global BioticEvent across the Paleocene–Eocene Boundary, in Sovre�mennye problemy geologii (Current Problems of Geology),Moscow: Nauka, 2004, pp. 493–531.

Gavrilov, Yu.O., Shcherbinina, E.A., and Oberhansli, H.,Paleocene–Eocene Boundary Events in the NortheasternPeri�Tethys, in Causes and Consequences of Globally WarmClimates in the Early Paleogene, Wings S.L. et al. Eds., Spec.Pap. Geol. Soc. Am., 2003, no. 369, pp. 147–168.

Gavrilov, Yu.O., Kodina, L.A., Lubchenko, I.Yu., andMuzylev, N.G., The Late Paleocene Anoxic Event in Epi�continental Seas of the Peri�Tethys and Formation ofSapropelite Unit: Sedimentology and Geochemistry, Litol.Polezn. Iskop., 1997, no. 5, pp. 492–517 [Lithol. Miner.Resour. (Engl. Transl.), vol. 32, no. 5, pp. 427–450].

Gibbs, S.J., Bown, P.R., Sessa, J.A., et al., NannoplanktonExtinction and Origination across the Paleocene–EoceneThermal Maximum, Science, 2006, vol. 314, pp. 1770–1773.

Giusberti, L., Rio, D., Agnini, C., et al., Mode and Tempoof the Paleocene–Eocene Thermal Maximum in anExpanded Section from the Venetian Pre�Alps, GSA Bull.,2007, vol. 119, nos. 3–4, pp. 391–412.

Gorbach, L.P., Stratigrafiya i fauna mollyuskov rannegopaleotsena Kryma (Stratigraphy and Malacofauna of theEarly Paleocene from Crimea), Moscow: Nedra, 1972.

Heilmann�Clausen, C., Dinoflagellate Stratigraphy of theUppermost Danian to Ypresian in the Viborg 1 Borehole,Central Jylland, Denmark, Pap. Geol. Surv. Denmark, Ser. A1985, no. 7, pp. 1–69.

Heilmann�Clausen, C., Review of Paleocene Dinoflagel�lates from the North Sea Region, in Proc. Meeting “Stratig�raphy of the Paleocene,” GFF, 1994, vol. 116, pp. 51–53.

Iakovleva, A.I., Brinkhuis, H., and Cavagnetto, C., LatePalaeocene–Early Eocene Dinoflagellate Cysts from theTurgay Strait, Kazakhstan: Correlations across Ancient

Seaways, Palaeogeogr. Palaeoclimatol. Palaeoecol, 2001,vol. 172, pp. 243–268.Iakovleva, A.I. and Heilmann�Clausen, C., Wilsonidiumpechoricum New Species – a New Dinoflagellate Specieswith Unusual Asymmetry from the Paleocene–EoceneTransition, J. Palaeontol., 2007, vol. 81, no. 5, pp. 1020–1030.Kennett, J.P. and Stott, L.D., Abrupt Sea�Level Warming,Paleoceanographic Changes and Benthic Extinctions at theEnd of Paleocene, Nature, 1991, vol. 353, pp. 225–229.Luterbacher, H.P., Ali, J.R., Brinkhuis, H., et al., ThePaleogene Period, in A Geologic Time Scale 2004, Gradstein F.M.,et al., Eds., Cambridge: Cambridge Univ. Press, 2004,pp. 384–408.Monechi, S. and Angori, E., Data Report: CalcareousNannofossil Biostratigraphy of the Paleocene–EoceneBoundary, Ocean Drilling Program Leg 208 Hole 1266C, inProc. Ocean Drilling Program. Scientific Results, vol. 208,Kroon, D., Zachos, J.C, and Richter, C., Eds., doi:10.2973/odp.proc.sr.208.203.2006.Mudge, D.C. and Bujak, J.P., Paleocene Biostratigraphyand Sequence Stratigraphy of the UK Central North Sea,Mar. Petrol. Geol., 1996, vol. 13, pp. 295–312.Muzylev, N.G., Anoxic Events of the Paleocene–MiddleEocene, in Ekosistemnye perestroiki i evolyutsiya biosfery(Ecosystem Turnovers and Evolution of Biosphere), Roza�nov, A.Yu. and Semikhatov, M.A., Eds., Moscow: Nedra,1994.Nemkov, G.I. and Barkhatova, N.N., Nummulity, assiliny ioperkuliny Kryma (Nummulites, Assilines and Operculinesof the Crimea), Moscow–Leningrad: Musei Karpinskogo,1961.Nunes, F. and Norris, R.D., Abrupt Reversal in OceanOverturning During the Paleocene–Eocene Warm Period,Nature, 2006, vol. 439, no. 5, pp. 60–63.Powell, A.J., Dinoflagellate Cysts of the Tertiary System, inA Stratigraphic Index of Dinoflagellate Cysts, British Micro�paleontol. Soc. Publ. Series, London: Chapman and Hall,1992, pp. 155–251.Powell, A.J., Brinkhuis, H., and Bujak, J.P., Upper Pale�ocene–Lower Eocene Dinoflagellate Cysts Sequence Bios�tratigraphy of Southeast England, Geol. Soc. Spec. Publ.,1996, no. 101, pp. 145–183.Shutskaya, E.K., Stratigrafiya, foraminifery i paleo�geografiya nizhnego paleogena Kryma, Predkavkaz’ya izapadnoi chasti Srednei Azii (Stratigraphy, Foraminifers andPaleogeography of the Lower Paleogene in Crimea, Ciscau�casia and Western Middle Asia), Moscow: Nedra, 1970.Sluijs, A., Pross, J., and Brinkhuis, H., From Greenhouseto Icehouse: Organic�Walled Dinoflagellate Cysts as Pale�oenvironmental Indicators in the Paleogene, Earth Sci.Rev., 2005, vol. 68, pp. 281–315.Sluijs, A., Bowen, G.J., Brinkhuis, H., et al., The Palae�ocene–Eocene Thermal Maximum Super Greenhouse:Biotic and Geochemical Signatures, Age Models andMechanisms of Global Change, in Deep�Time Perspectiveson Climate Change: Marrying the Signal from Computer Mod�els and Biological Proxies, Williams M., Haywood A.M.,Gregory F.J., Schmidt D.N., Eds., Spec. Publ. Micropalae�ontol. Soc. London, 2007, pp. 323–349.Speijer, R.P. and Wagner, T., Sea�Level Changes and BlackShales Associated with the Late Paleocene Thermal Maxi�



mum: Organic�Geochemical and Micropaleontologic Evi�dence from the Southern Tethyan Margin (Egypt–Israel),Spec. Pap. Geol. Soc. Am., 2002, vol. 356, pp. 533–549.Stratigrafiya (shel’f i poberezh’ya Chernogo morya) (Stratig�raphy of the Black Sea Shelf and Coast), Kiev: Naukovadumka, 1984.Stupin, S.I. and Muzylev, N.G., The Late Paleocene Eco�logic Crisis in Epicontinental Basins of the Eastern Peri�Tethys: Microbiota and Accumulation Conditions ofSapropelic Bed, Stratigr. Geol. Korrelyatsiya, 2001, vol. 9,no. 5, pp. 501–507 [Stratigr. Geol. Correlation (Engl.Transl.), vol. 9, no. 5, pp. 501–507].Thomas, E., The Biogeography of the Late PaleoceneBenthic Foraminiferal Extinction, in Late Paleocene–EarlyEocene Biotic and Climatic Events in the Marine and Terres�trial Records, Aubry M.P., et al., Eds., New York: ColumbiaUniv. Press, 1998, pp. 214–243.Tremolada, F. and Bralower, T.J., Nannofossil AssemblageFluctuations During the Paleocene–Eocene Thermal

Maximum at Sites 213 (Indian Ocean) and 401 (NorthAtlantic Ocean): Paleoceanographic Implications, Mar.Micropaleontol., 2004, no. 52, pp. 107–116.

Tripati, A. and Elderfield, H., Deep�Sea Temperature andCirculation Changes at the Paleocene–Eocene ThermalMaximum, Science, 2005, vol. 308, no. 5730, pp. 1894–1898.

Vasil’eva, O.N., Dinocysts from the Paleocene–EoceneBoundary Deposits in the Southern Trans�Urals, in Ezhe�godnik�1999 (Yearbook 1999), Yekaterinburg: Inst. Geol.Geokhim., 2000, pp. 11–16.

Wing, S.L., Harrington, G.J., Smith, F.A., et al., TransientFloral Change and Rapid Global Warming at the Pale�ocene–Eocene Boundary, Science, 2005, vol. 310, pp. 993–996.

Zachos, J., Pagani, M., and Sloan, L.C., Trends, Rhythms,and Aberrations in Global Climate 65 Ma to Present, Sci�ence, 2001, vol. 292, pp. 686–693.

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