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MASARYKOVA UNIVERZITA
PŘÍRODOVĚDECKÁ FAKULTA
Ústav geologických věd
Palynologie a její využití pro interpretace
přírodního prostředí a jeho změn
Habilitační práce
Nela Doláková
Brno 2018
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Bibliografický záznam
Habilitační práce
Autor: RNDr. Doláková Nela, CSc.
Přírodovědecká fakulta, Masarykova univerzita
Ústav geologických věd
Název: Palynologie a její využití pro interpretace přírodního prostředí a jeho
. změn.
Počet stran: 296
Klíčová slova: Palynologie, metodika, paleoekologie, neogén, pleistocén, holocén,
marinní sedimenty, jeskynní sedimenty, archeologické lokality
Nela Doláková, Masarykova univerzita, 2018
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Bibliographic Entry
Author: RNDr. Doláková Nela, CSc.
Faculty of Science, Masaryk University
Institute of Geological Sciences geologických věd
Title: Palynology and its application for the interpretations of natural
environments and their changes
Number of pages: 296
Keywords: palynology, methodology, palaeoecology, Neogene, Pleistocene, Holocene,
marine sediments, cave sediments, archaeological localities
Nela Doláková, Masaryk University, 2018
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Poděkování:
Chtěla bych poděkovat všem kolegům i své rodině za trpělivost, pomoc a morální podporu
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Abstrakt
Předkládaná práce je zaměřena na problematiku (paleo)palynologie a jejího využití pro
interpretace přírodního prostředí a jeho změn.
Informace získané ze studia palynomorf v horninách můžeme využít zejména pro
rekonstrukce vegetačního pokryvu v minulosti a jeho změn, interpretace klimatu,
paleogeografie nebo stratigrafie. Publikované práce jsou zaměřeny nejen na studium vegetace
v terestrických sedimentech, ale i interpretace zachovaných pylových spekter ze sedimentů
mořských. V práci je využito 17 autorských publikací.
Práce je rozdělena do několika tematických okruhů. První okruh zahrnuje metodické přístupy
a obecnou problematiku palynologie. Náplní této kapitoly jsou principy zachování a degradace
palynomorf v horninovém prostředí a s těmito procesy spojená problematika redepozic.
V kapitole je rovněž přiblížena tématika přípravy vzorků, principů mikroskopického
pozorování, základních příkladů popisu zrn a kvantitativního vyhodnocování pylových spekter.
V kapitole jsou využity nejen teoretické principy zobecněné v literatuře, ale i vlastní zkušenosti
autorky. Druhý okruh komentovaných palynologických studií se týká interpretace vegetace
v sedimentech neogenního stáří. Kapitola je rozdělena do dvou částí. První část je zaměřena na
interpretace vegetace a jejích změn v čase, které se odrážejí zejména ve vývoji klimatu. Ve
druhé části jsou výsledky palynologie v koordinaci s dalšími paleontologickými,
sedimentologickými a geochemickými metodami využity k řešení širší geologické
problematiky, zejména charakteru mořských pánví, jejich okrajů a vzájemného ovlivňování
s prostředím kontinentu. Kapitola - Palynologie vybraných lokalit kvartéru je věnována
problematice interpretace vegetačních poměrů ze studia jeskynních sedimentů a
archeologických lokalit. Největším problémem interpretace pylových spekter z jeskynních
sedimetů je jejich transport do míst ukládání spojený s možností sekundární selekce zrn.
Problematika studia palynologie z archeologických lokalit spojuje nejen změny klimatických
charakteristik, ale rovněž změny vegetace v důsledku lidských aktivit.
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Abstract
The proposed paper is focused on the (paleo) palynology and its use for the interpretation of
natural environment and its changes.
The information obtained from the palynomorph study in the rocks can be used especially for
the reconstruction of the vegetation cover in the past and its changes, the interpretation of
climate and paleogeography or the stratigraphical determination. The published works are
focused not only on the study of vegetation in terrestrial sediments, but also on the
interpretations of preserved pollen specter from marine sediments. The work uses 18 author's
publications.
The thesis is divided into several thematic scopes. The first round includes methodological
approaches and the general problems of palynology. The contents of this chapter are the
principles of conservation and degradation of palynomorph in the rock environment and these
processes are connected with problems of redepositions. The chapter also deals with the topic
of sample preparations, principles of microscopic observations, basic examples of grain
descriptions and quantitative evaluation of pollen spectra. The chapter uses not only the
theoretical principles generalized in the literature, but also the author's own experience.
The second round of commented palynological studies refers to the interpretation of vegetation
in sediments of the Neogene ages. The chapter is divided into two parts. The first part is focused
on the interpretation of vegetation and its changes in time, which are reflected mainly in the
climate. In the second part, the results of palynology, in coordination with other
palaeontological, sedimentological and geochemical methods, are used to resolve broader
geological problems, especially the character of the sea basins, their margins and mutual
interactions with terrestrial environment.
The chapter on palynology of selected Quaternary localities is devoted to the problem of
interpretation of vegetation conditions from the study of cave sediments and archaeological
sites. The main problem of interpretation of pollen spectra from cave sediments is their transport
to sedimentation places associated with the possibility of secondary grain selection. The scope
of palynological study from archaeological sites combines not only changes in climatic
characteristics but also changes in vegetation due to human activities.
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Obsah:
1. Úvod…………………………………………………………………………. 8
2. Metodické přístupy a obecná problematika palynologie……….……….. 10
2a) Tafonomické procesy - zachování x degradace palynomorf………………....... 10
2b) Problematika redepozic…………………………………….………………….. 12
2c) Metody studia ………………………………………….……………………… 14
2ca) Příprava vzorků……………………..………………………….….………… 14
2cb) Mikroskopické techniky, základní principy popisu zrn …………...…...….. 16
2cc) Kvantitativní vyhodnocení - tvorba pylových diagramů……………………. 20
3. Palynologie neogénu …………………………………………………….….. 21
3a) Interpretace vegetace a klimatických změn…………………………………… 25
3ab) Charakteristika vegetace v jednotlivých časových úsecích …....………….. 31
3b) Širší problematika neogénu v koordinaci palynologie a dalších
paleontologických, sedimentologických a geochemických metod výzkumu…
38
4. Palynologie vybraných lokalit kvartéru………..………………………...… 41
4a) Jeskynní sedimenty………………………………………………………….... 42
4b) Archeologické lokality – vliv člověka na prostředí …………………………. 47
5. Použitá literatura…………………………………………………….…...... 53
6. Přílohy……………………………………………………………………… 67
6a) Seznam přiložených publikací použitých v textu jednotlivých kapitol……… 67
6b) Přiložené publikace ..……………………………………………………...... 69
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1. Úvod
Předkládaná práce je zaměřena na problematiku paleopalynologie a jejího využití pro
interpretace přírodních procesů a jejich změn.
Palynologie je disciplína, která zkoumá zejména pylová zrna a spory rostlin. Patří sem ovšem i
studium dalších mikroobjektů s tzv. acidorezistentními obaly, které se většinou shrnují do
skupiny tzv. nepylových objektů. Jedná se např. o mořský i sladkovodní fytoplankton,
mikroskopické zbytky hub, živočišné parazity, zbytky srsti nebo těl hmyzu apod. Obecně se
pro všechny tyto objekty používá název palynomorfy.
V rámci geologie se často hovoří o paleopalynologii, kdy se studují palynomorfy, které
prošly procesem fosilizace.
Rozvoj oboru je umožněn díky velké schopnosti zachování palynomorf, jejich
morfologické rozmanitosti a snadnému transportu (větrem, vodou a živočichy). Můžeme je
nalézt téměř ve všech typech sedimentárních hornin, a dokonce i v horninách slabě
metamorfovaných. Palynomorfy můžeme studovat jak v sedimentech terestrických, tak i
marinních, což představuje jednu z mála možností pro korelace těchto vývojů.
Informace získané ze studia palynomorf v horninách můžeme využít zejména pro
rekonstrukce vegetačního pokryvu v minulosti a jeho změn, interpretace klimatu i
paleogeografie nebo stanovení stratigrafie. Interpretovat můžeme nejen charakter
suchozemského prostředí, ale i některé faktory prostředí mořského (např. hloubky, vzdálenost
od pobřeží, oxidačně-redukční procesy) a jejich vzájemné ovlivňování – morfologie pobřeží,
změny salinity, rozšiřování a ústup mořských pánví.
Jako u každé metody existuje celá řada omezení vypovídací hodnoty studovaných
palynospekter. Je proto velmi vhodné kombinovat znalosti z palynologie a dalších metod (např.
paleontologie, sedimentologie, geochemie atd.), které při vzájemném doplnění poznatků
umožní co nejpřesnější přiblížení se k poznání obrazu minulosti a jeho vývoje.
Předkládaná práce shrnuje výsledky studia palynologie v sedimentech poměrně velkého
časového rozsahu. Zahrnuje období v průběhu posledních 22 milionů let (neogén, pleistocén a
holocén – více než 80 lokalit). Každý z těchto časových úseků se vyznačuje specifickým
vývojem sedimentace a rozličnými ekologickými a tafonomickými podmínkami. Proto byla
následující práce rozdělena do několika oddílů, které se po úvodním shrnutí obecné
problematiky soustřeďují na dosažené výsledky studia v jednotlivých obdobích. V práci je
využito a komentováno celkem 17 autorských publikací. Z důvodů kompaktnosti zvolených
tématických celků bylo použito nejen publikací citovaných v databázích WOS a Scopus, (15),
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ale rovněž 2 prací v recenzovaných časopisech a 1 kapitoly v monografii. Práce, jejichž
výsledky se týkají nejen konkrétních palynologických interpretací, ale i zobecňující metodické
problematiky, jsou v textu využity v různých kapitolách. Výsledky některých dalších prací jsou
pouze citovvány a v krátkosti zmíněny, nejsou však zahrnuty do výčtu komentovaných prací.
Nejzajímavější výsledky týkající se zejména klimatických a paleogeografických změn
nebo dopadů lidské činnosti na vegetaci jsou využívány i ve výuce pro studenty geologických
a archeologických oborů MU.
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2. Metodické přístupy a obecná problematika palynologie
2a) Tafonomické procesy - zachování x degradace palynomorf
Vysoká fosilizační schopnost palynomorf je umožněna díky složení jejich stěn,
tvořených tzv. sporopolleninem. Je to organická polymerní látka příbuzná chitinu, která je
odolná vůči působení kyselin – proto je označována jako acidorezistentní. Do jisté míry
odolává i zvýšené teplotě a tlaku až do účinků slabší metamorfózy (Brooks1971). Pylová zrna
a spory se proto vyskytují v sedimentárních, ale i slabě metamorfovaných horninách.
Nejlépe se palynomorfy zachovávají ve vlhkém kyselém prostředí (např. rašeliny), méně
v jemně klastických uloženinách mořského, říčního, deltového, jezerního nebo jeskynního
prostředí (Moore, Webb, Collinson 1991) a rovněž v některých typech archeologických
objektů.
Studium palynomorf v sedimentárních profilech má výhodu zejména v dlouhém časovém
záznamu, kdy můžeme sledovat změny jednotlivých faktorů prostředí (např. teplota, vlhkost,
povaha substrátu), ale i vymírání, migrace, nebo objevování se nových taxonů. Nevýhodou je
(obdobně jako u každého fosilního záznamu) převážně selektivní zachování – výsledným
obrazem je tzv. oryktocenóza.
Oproti makrofloristickým nálezům má palynologie několik výhod a naopak nevýhod.
Zatímco rostlinné makrozbytky lze většinou determinovat do úrovně rodu i druhu, palynomorfy
jsou v mnohých případech určitelné maximálně do úrovně rodu nebo jen čeledi, ve starších
obdobích dokonce pouze jako morfotypy. U rostlinných makrozbytků (zejména listů a kutikul)
lze i u vymřelých taxonů posuzovat taxonomické zařazení na základě srovnávací anatomie a
morfologie. Tato možnost je u palynomorf omezená. Na druhou stranu se makrozbytky
uchovávají většinou jen v sedimentech z mokřadních prostředí s nízkou dynamikou vody
(kromě archeologických situací), kde jsou zachovány fosílie především z těchto ekologických
prostředí a jejich bezprostředního okolí. Taxony a společenstva vyskytující se ve větší
vzdálenosti od těchto k fosilizaci vhodných míst mají jen velmi malou možnost uchování (např.
taxony sušších nebo horských stanovišť). V oryktocenózach palynomorf se nám díky značné
transportní schopnosti dochovávají fosílie rostlin i z prostředí nepříznivých pro zachování
fosilních zbytků. Získáváme tedy komplexnější obraz širšího paleoprostředí. Při
palynologických interpretacích je ovšem třeba obezřetně řešit i problematiku rozdílné pylové
produkce a transportní schopnosti palynomorf jednotlivých rostlin (rostliny hmyzosnubné x
větrosnubné), ve fosilních asociacích navíc rozdílnou fosilizační schopnost zrn.
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Během transportu a sedimentace palynomorf může vlivem tafonomických procesů
docházet k jejich degradaci až zničení, případně ke druhotné akumulaci odolnějších zrn (časté
např. u jeskynních a mořských prostředí, viz následující kapitoly). Na degradaci se podílí jak
mechanické, tak chemické či biologické, především bakteriální vlivy (např. Havinga, 1964,
1967, 1971). Výsledná oryktocenóza potom může poskytovat značně zkreslený obraz původní
vegetace.
Velmi nepříznivě se na zachování palynomorf podílí oxidace. Zvýšená oxidace se
projevuje v pórovitých – hrubě klastických horninách (zde navíc silné mechanické poškození
vlivem transportu klastik - písky, štěrky). V dobře prokysličeném prostředí vzniká i většina
kalcitických sedimentárních hornin (vápence, travertin). Oxidační procesy probíhají i
v některých typech sedimentů, jejichž vznik je klimaticky podmíněný (jíly pestrých barev,
kaolinit). V těchto typech hornin je možnost zachování palynomorf velmi nepravděpodobná.
V závislosti na chemickém složení a tloušťce sporopylových obalů podléhají
palynomorfy různých rostlin rozkladu vlivem oxidace různou rychlostí (např. pylová zrna
čeledi Lauraceae se v sedimentech nedochovávají). Experimentálním studiem odolnosti
pylových zrn vybraných rostlinných taxonů v závislosti na zvyšující se intenzitě oxidace se
zabývali např. Kwiatowski , Lubliner-Mianowska (1957), Havinga (1971), Hopkins, Mc Carthy
(2002)
V souvislosti se stupněm zachování palynomorf byly detekovány změny oxidačněredukčních
podmínek i v mořských miocenních sedimentech z karpatské předhlubně na Moravě
(Doláková et al. 2014, Holcová et al. 2015, Nehyba et al. 2016). Zde bylo v několika vrtech
a lokalitách pozorováno periodické střídání palynospekter s pestrými společenstvy a
palynospekter, kde převládala pylová zrna jednoho typu. Jednalo se téměř výhradně o pylová
zrna konifer (Pinus, Cathaya, Picea, Cedrus). Tato zrna se sice díky své schopnosti dalekého
doletu mohou akumulovat i v mořských oblastech daleko od břehu, ale podle výsledků studia
dalších metod (geochemie, paleontologie, sedimentologie) periodické změny prostoru mořské
pánve nebyly předpokládány (viz kap. 3.).
V mořských prostředích s výrazným nedostatkem kyslíku (hypoxické – anoxické)
dochází v sedimentu při redukčních procesech k tvorbě mikrokrystalů pyritu. Mohou se tvořit
i uvnitř pylových zrn. Po chemické maceraci jsou patrné jako typické drobné krychlovité
dutiny, které zůstanou na zrnech viditelné. Mohou být vyvinuty v takovém množství, že způsobí
změny morfologie až destrukci zrn (Doláková, Slamková 2003, Kováčová et al., 2011).
Z charakteru pylových asociací i morfologických změn na pylových zrnech a sporách lze
tedy usuzovat i na změny oxido-redukčního potenciálu sedimentačního prostředí.
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Degradace palynomorf mechanickými vlivy je způsobena interakcemi zrn s částicemi
sedimentu během intenzivního transportu do místa ukládání. Během těchto procesů se
dochovávají jen nejmenší nebo morfologicky nejkompaktnější zrna. Tato jsou potom ve
výsledné oryktocenóze druhotně nakumulovaná a pro případné interpretace vegetace
nadhodnocená. Proces je typický zejména pro uloženiny v jeskynním prostředí a rovněž pro
prostředí chladného klimatu s převažující sedimentací klastik, např. spraší – Asteroideae –
hromadění v chladných obdobích (Doláková 2007, 2014).
Biologické vlivy se uplatňují především v sedimentačních prostředích vhodných pro
činnost mikroorganizmů (např. bakterií). Tento proces je nejintenzivnější při půdotvorných
procesech. V půdních horizontech dochází většinou ke kompletní ztrátě palynologických dat.
Při studiu palynologie je vhodné rovněž zaznamenávat a pokud možno determinovat tzv.
nepylové objekty. Jejich vypovídací schopnost je zejména v oblasti ekologie a paleoekologie.
Jedná se např. o cysty nebo cenobia řas (mořské, sladkovodní, půdní – chladnomilné x
teplomilné, oligotrofní x eutrofní), mikroskopické zbytky hub (např. dřevokazné, mykorhizní,
koprofilní), organické výstelky foraminifer (tzv. tapeta), obrněnky, vajíčka živočichů (např.
cizopasných).
2b) Problematika redepozic
V průběhu opakujícího se procesu sedimentace a zvětrávání nejsou vyloučeny redepozice
(přemístění) palynomorf ze sedimentů vzniklých ve starších časových obdobích, do sedimentů
mladších, a tím smíšení zrn z různých stratigrafických úrovní. Existence a množství
redeponovaných palynomorf závisí na prostředí a typu sedimentace. Nejčastější jsou
v sedimentech, které vznikly erozí, transportem a opětovným ukládáním klastik (typicky
jeskynní a mělkomořské prostředí). Pokud nelze redeponovaná zrna odlišit od zrn in situ, může
docházet k chybným interpretacím stáří i prostředí. Z geologického hlediska mohou být však
takovéto redeponované palynomorfy jediným důkazem o původní existenci sedimentů, které
byly z daného území kompletně denudovány. Příkladem jsou pylová zrna skupiny Normapolles
(skupina vymřelých krytosemenných, rozšířených zejména během křídy), která jsou nalézána
v neogenních mořských sedimentech karpatské předhlubně a svědčí o mnohem větším rozšíření
křídových sedimentů, než známe z území jižní Moravy dnes (Doláková, Slamková 2003).
Jednoznačné odlišení redeponovaných palynomorf je možné pouze v případě
determinace typů, které jsou v časovém období reprezentovaném studovaným sedimentem
vymřelé nebo pocházejí z prostředí, které neodpovídá procesu sedimentace (např. mořská
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dinoflagellata v sedimentech spraší). V jiných případech mají redeponovaná zrna odlišný
stupeň fosilizace reprezentovaný např. změnou barvy nebo tloušťkou pylových stěn.
Často je však rozlišení redeponovaných zrn velmi obtížné, zejména u smíšení sedimentů
z časově blízkých období, která obsahují obdobné typy rostlin - např. u mořských sedimentů
jednotlivých miocenních stupňů, odkud jsou běžně známé i další redeponované fosílie (např.
foraminifery a vápnitý nanoplankton).
Složitou problematiku představují redepozice v jeskynních sedimentech (Doláková
2007). Většina jeskynních sedimentů z moravských krasových oblastí je kvartérního stáří.
Mnoho rostlin, jejichž pylová zrna nebo spory v těchto sedimentech můžeme najít, se na našem
území vyskytuje od neogénu do současnosti - příkladem jsou stromy jako borovice, jilm, olše,
dub, lípa, ale i byliny jako trávy, složnokvěté nebo i některé druhy kapradin a další. Mnohé
z nich jsou přitom velmi důležité pro stanovení jednotlivých teplotních fází kvartéru
(klimatostratigrafie). Tento fakt velmi ztěžuje přesné a jednoznačné stanovení stáří sedimentů,
kdy nemůžeme stoprocentně odlišit tercierní redepozice a sediment se může jevit jako vzniklý
v daleko teplejším období. (viz kap. 4a).
Jednou z metod, které mohou přispět k rozlišení redeponovaných pylových zrn,
představuje studium ve fluorescenčním mikroskopu. Metody využití UV fluorescence pro
determinaci různě starých palynomorf rozpracoval Van Gijzel (1967a ,b, 1971, 1975, 1978) či
Yeloff, Hunt (2005). Organický materiál stěny palynomorf vykazuje při působení UV záření
autofluorescenci. Spektrum a intenzita fluorescenčních barev závisí na chemickém složení
organických stěn. Toto chemické složení se mění s geologickým stářím a při procesech
probíhajících v sedimetech – uhelnatění nebo zvětrávání spojené s oxidací a bioerozí. Při
redepozicích proto dochází ke změně intenzity a barvy fluorescenčního spektra studovaných
taxonů. Praktickým využitím těchto metod na studiu zejména jeskynních, ale i mořských
sedimentů se zabývá práce Doláková, Burešová (2007). Pylová zrna různých rostlinných
taxonů vykazují odlišnou úroveň fluorescence v závislosti na chemickém složení a tloušťce
exiny (např. pylová zrna trav jeví nízkou intenzitu fluorescence). Při studiu celého pylového
spektra nelze proto z úrovně barev říci, zda některé taxony jsou redeponované a jiné in situ. Pro
běžné využití tohoto jevu bez dalších složitých měřících přístrojů je nutné zaměřit se na studium
jednoho taxonu (odlišná úroveň fluorescence u zrn v jednom vzorku). Pokud ovšem daný taxon
má v UV světle velmi tmavé barvy s velmi nízkou intenzitou, tzn. téměř nesvítí, jedná se velmi
pravděpodobně o redepozici. Odolné vůči chemickým procesům jsou organické stěny
mikroskopických řas a jejichy cysty – vykazují intenzívní fluorescenci. Naopak zbytky
14
živočišných těl – např. různé typy vajíček, jeví tmavé a málo intenzivní fluorescenční barvy
(Doláková, Burešová 2007).
Studium fluorescence bylo využito pro detekování případných redepozic u pylových zrn
rodu Carpinus (habr) z holocenních sedimentů na Pohansku (Doláková et al., 2010). Na
základě měření stáří metodou 14
C byla pylová zrna habru zjištěna v sedimentech nejvyššího
stáří na území ČR (8 000 let). Na základě studia fluorescence byly redepozice z tercierních
sedimentů v podloží vyloučeny.
Značnou odolnost vůči oxidačním procesům spojeným se zvětráváním, a tudíž i
redepozicemi, mají pylová zrna konifer (Kwaitkowski, Lubliner- Mianowska 1957, Havinga
1964, 1967, Brooks 1971, Hopkins, Mc Carthy 2002). Poměr redeponovaných a in situ zrn
konifer s létacími vaky byl studován za pomoci UV-fluorescence ve vzorcích z vrtu Oslavany1
(Nehyba et al. 2016), kde tato zrna tvořila většinu pylového spektra. Pouze malá část zrn jevila
odlišný, velmi nízký stupeň fluorescence. Proto byla většina asociace z tohoto hlediska
považována za autochtonní a vysvětlení nadměrného množství konifer mělo jiné příčiny (viz
kap. 3a, b).
Dalším tafonomickým jevem je uchování pylových zrn jednoho druhu ve shlucích. Tento
jev ukazuje na ukládání v blízkosti původní rostliny, která pyl produkovala. Z geologického
hlediska se tedy jedná o depozici in situ, bez výrazného ovlivnění transportem (nízká dynamika
vody). Pozorované shluky byly v tercierních sedimentech typické např. pro sedimentaci v
pobřežních bažinách, deltových prostředích nebo tzv. marších (Myricaceae, Chenopodiaceae,
Caryophyllaceae, Oleaceae, Onagraceae, Platanus) (Doláková et al. 1999, Doláková 2004,
Kováčová et al. 2011).
V kvarterních sedimentech je tento jev zajímavý i z archeologického hlediska, např. v
profilu holocenními sedimenty na lokalitě Nad Velkým Jejkalem (Národní park Podyjí) byla
identifikována pylová zrna obilí typu pšenice (Triticum), která zůstala zachována v
nerozpadených prašnících. Tento nález svědčí o pěstování velmi blízko místa uložení. Vzorek
z této hloubky (85-94 cm), byl datován radiokarbonovou metodou jako doba římská a období
stěhování národů – tyto nálezy mohou indikovat existenci blízkého lidského sídliště, které
dosud nebylo archeologicky prokázáno (Šušolová et al. 2016).
15
2c) Metody studia
2ca) Příprava vzorků
Vzhledem k malým rozměrům palynomorf a jejich rozptýlení v sedimentech je nutné je
z těchto hornin separovat a zkoncentrovat. Využívá se různých postupů macerace, kdy je třeba
odstranit co nejvíce anorganických minerálních součástí a organickou hmotu koncentrovat a
prosvětlit. Macerace představuje kombinace působení chemických procesů a mechanické
přípravy založené na rozmělňování, sítování a hmotnostní separaci (využití těžkých kapalin).
Kombinace jednotlivých procesů se přizpůsobuje typu horniny. Nejčastěji používanou metodou
jetzv. Erdtmanova metoda acetolýzy (Erdtman1960). Soubor postupů podle různých autorů
s přizpůsobeními pro jednotlivé typy hornin shrnuly např. Pacltová 1963, Gabrielová 1986.
Alternativní metody macerací, kde jde zejména o vyloučení práce s kyselinou fluorovodíkovou,
publikovali Riding, Kyffin-Hughes (2006). V naší praxi se prozatím tyto metody neosvědčily.
Nejpoužívanější standardní palynologická macerace pro sedimenty terciéru a kvartéru je
založená na těchto principech:
1. Odstranění karbonátů pomocí kyseliny chlorovodíkové (HCl)
2. Odstranění silikátů - kyselina fluorovodíková (HF)
3. Odstranění vytvořených fluoridových gelů (HCl)
4. Hmotnostní separace minerálního rezidua a zkoncentrování pylonosné frakce
za pomoci těžké kapaliny (ZnCl2, BrCl2) - hustota = 2g/cm3
5a) U uhelných sedimentů je třeba ještě provést zesvětlení zuhelnatělých palynomorf
pomocí oxidace: HNO3, KOH
5b) U palynomorf z mladých – holocenních sedimentů, kde ještě nedošlo ke kompletní
fosilizaci, je třeba prosvětlit palynomorfy a jejich organický obsah pomocí tzv. acetolýzy H2SO4
+ acetanhydrid kyseliny octové (CH3COOH). Vynechání acetolýzy u pleistocenních a
starších sedimentů naopak umožní odlišení kontaminace recentními pylovými zrny a sporami
např. při průsaku srážkové vody po puklinách nebo kořenech rostlin (zůstane zachována
cytoplazma).
Hmotnost zpracovávaných vzorků a volba metody macerace závisí na typu horniny.
Odebírá se od několika gramů až po půl kilogramu. Nejvyšší koncetrace a nejlepší zachování
palynomorf lze očekávat v sedimentech, které vznikaly v prostředí bez přístupu kyslíku nebo
druhotných oxidačních procesů a snížené dynamiky, kde dochází k hromadění organické hmoty
(např. rašeliny, sedimenty jezerní, deltové, pobřežních lagun a bažin, slepých ramen řek apod.).
16
1. rašeliny - pomalé ukládání sedimentu, nízká nebo žádná minerální příměs -vysoká
koncentrace palynomorf - malé vzorky (cca 20g);
2. sediment s velkou příměsí minerálního materiálu – vysoká rychlost akumulace
sedimentu, nízká koncentrace palynomorf - větší vzorky (500g), např. povodňové hlíny
nebo spraše
Při uchovávání a dalším studiu vymacerovaných palynomorf je nutné zamezit další
oxidaci. Výsledný macerát se proto uchovává ve směsi glycerínu, ethanolu a vody.
Mikroskopická pozorování lze provádět přímo v tomto tekutém médiu – pylové zrno je
možné posunovat a otáčet. Potřebujeme-li preparáty trvalé (uchování preparátu po delší čas),
používá se většinou tzv. glycerinová želatina nebo kanadský balzám.
2cb) Mikroskopické techniky, základní principy popisu zrn
Palynomorfy se převážně determinují v biologických prosvěcovacích mikroskopech na
základě jejich morfologické rozmanitosti. Běžně je používáno zvětšení 200x, 400x a 1000x imerzní
objektiv). Výhodou biologických mikroskopů je posuvný stolek, kde je možné za
pomoci dvou na sobě kolmých měřítek zaznamenat přesnou polohu zrna v preparátu.
K popisu a determinaci pylových zrn byla vytvořena ustálená terminologie založená
především na morfologii zrn. Nejdůležitější znaky souvisejí s postavením zrna v tetrádě,
vnějším tvarem a velikostí, skulptuře a struktuře stěny, stavbou germinálního aparátu. První
souborný atlas morfologických znaků sestavil Erdtman (1957). Nejnovější shrnutí
palynologické termilogie představuje práce Punt et al. (2007). Příklady morfologických znaků
jsou znázorněny na obr. 1- 5.
a) b) Caluna sp.
Obr 1. a) Základní tvary tetrád spor a pylových zrn (upraveno Punt et al. 2007)
b) tetráda pylového zrna rodu Calluna (foto autorka)
17
a) b)
Obr. 2. Rozdíly velikostí pylových zrn: a) pylové zrno rodu Abies, b) pylové zrn rodu Castanea
(foto autorka)
a)
b) Retikulátní Potamogeton klavátní Ilex
Obr. 3. a) Členění vnějšího obalu pylových zrn a spor
b) Příklady skulptury a ornamentace povrchu zrn: retikulátní - Potamogeton
klavátní - Ilex (upraveno Punt et al. 2007, foto autorka)
exina
intina
tectum
sloupková vrstva
bazální vrstva
Abies
Castanea
20µm
18
a) pór Poa Engelhardia Carya
b ) anulus Betula
Obr. 4. Příklady hlavních typů apertur (klíčních štěrbin), jejich umístění a kombinací:
a) Pór - pylová zrna s 1 pórem - Poa, 3 póry v různém unítění - Engelhardia, Carya
b) Příklad vnitřní stavby póru - anulus, pylové zrno s anulem - Betula
(upraveno Punt et al. 2007, foto autorka)
a) Kolpa Quercus Galium
b) Kombinace póru a kolpy Cornus – Mastixia Sapotaceae
Obr. 5 Příklady hlavních typů apertur (klíčních štěrbin), jejich umístění a kombinací:
a) Kolpa - zrno se 3 kolpami – Quercus, se 6 kolpami – Galium
b) Kombinace póru a kolpy: 3x - Cornus - Mastixia typ, 4x Sapotaceae
(upraveno podle Punt et al. 2007, foto autorka)
19
U současných taxonů rostlin jsou pylová zrna a spory dostatečně prostudované k tomu,
aby se u jmen palynomorf mohlo využít systematické jméno odpovídajícího rostlinného
taxonu.
U fosilních palynomorf bohužel v mnohých případech konkrétní botanickou
příslušnost determinovaného taxonu stanovit nelze, protože část taxonů je vymřelých,
některé taxony exotické nejsou z hlediska pylové morfologie dostatečně prostudovány nebo
některé fosilní taxony mají vyšší morfologickou variabilitu, než je známa u taxonů
recentních.
Proto byla pro účely determinace fosilních palynomorf vytvořena umělá systematika,
kde jsou názvy taxonů založené na morfologických znacích pylových zrn a spor. Tato
systematika je využívána zejména ve starších pracích (např. Krutzsch 1966, 1971, Pacltová
1958, Nagy 1969, 1985, Planderová 1971, 1990). Morfotaxony se používaly i u názvů zrn,
jejichž botanická příslušnost byla nesporná. Část palynologů se naopak snaží pojmenovávat
pylové taxony podle botanické příslušnosti, zejména na základě detailnější morfologie
založené na studiu v elektronových mikroskopech (např. Zetter 1989, www.paldat.org).
V současné době se část specialistů palynologů snaží držet principu priority a používat pro
pylová zrna a spory modifikované názvy morfotaxonů (např. Stuchlik et al. 1994, 2014), i
když je botanická souvislost s originálními rostlinami jasná. Jako příklad můžeme uvést
různá synonyma pro pylová zrna rodu Juglans (ořešák): Multiporopollenites maculosus
(Potonié) Thomson & Pflug, Juglans Linne, Jugnlanspollenites juglandoides (KohlmanAdamska).
V detailních systematických pracích mívají proto pylové taxony velmi rozsáhlou
část synonymiky.
Z důvodů přesnější taxonomické determinace některých sporných palynomorf se
v poslední době často studují objekty v elektronovém mikroskopu. Využívá se zejména pro
determinaci a stanovení příbuznosti taxonů a hledání recentních ekvivalentů dosud
neurčených tercierních taxonů na základě morfologie povrchu pylových zrn a spor. Tyto
morfologické prvky nebývají v mikroskopech světelných zřetelné. Terminologii využívanou
pro morfologické charakteristiky při studiu v elektronovém mikroskopu (SEM) sestavil
kolektiv autorů Halbitter et al. (2006) v rámci databáze PALDAT (www.paldat.org). Je
ovšem nutné využít kombinaci studia daného objektu v prosvěcovacím mikroskopu (pohled
dovnitř zrna a vnitřní stavba terminálního aparátu) a téhož objektu v SEM (podrobná vnější
morfologie bez vnitřního pohledu). Metodiku tohoto studia rozpracoval Zetter (1989). Tato
metoda však vyžaduje speciální optické mikroskopy, které mají přídavnou optiku pro
20
vytvoření stranově nepřevráceného obrazu. Navíc je zde nutné objekty dále přenášet na
vodivé měděné terče, což neumožňuje opětovné pozorování ve světelném mikroskopu.
Na ÚGV PřF MU je k dispozici elektronový mikroskop JEOL JSM – 649 OLV, který
umožňuje vkládat studované objekty přímo na podložním skle. Pro palynologická studia
jsme s kolegyní M. Kováčovou (UK Bratislava) rozpracovaly metodiku přizpůsobenou
tomuto elektronovému mikroskopu s možností zpětného prohlížení v mikroskopu optickém
(Doláková, Kováčová připraveno do tisku).
Studia palynomorf založená na kombinované metodě LM/SEM byla využita zejména
pro detreminace palynomorf neogenního stáří. Poprvé bylo použito na zařazení morfotaxonu
Monocirculipollis Krutzsch a jeho přiřazení k čeledi Caryophyllaceae (Doláková 2004).
Metod LM/SEM bylo použito i pro řešení identifikace problematických morfotaxonů čeledi
Fagaceae: Tricolpopollenites liblarensis, Tricolporopollenites cingulum oviformis. Na
základě studia v SEM bylo zjištěno, že tato v optickém mikroskopu téměř identická zrna
mohou mít různou botanickou příslušnost – Castanea x Castanopsis x Trigonobalanopsis.
Jejich rozlišení je možné na základě morfologie povrchu zrn (Dolákova et al 2011).
Upřesnění identifikace na základě LM/SEM bylo provedeno i u rodů: Quercus, Platanus,
Tamarix, zástupců čeledí Caryophyllaceae a Rutaceae, rozlišení rodů Pinus x Cathaya.
Identifikace byla prováděna na základě srovnání s literárními údaji a rovněž na základě
studia recentních pylových zrn - Castanea sativa, Tamarix gallica, Citrus limon (Doláková
et al. 2011, Kováčová et al. 2011)
2cc) Kvantitativní vyhodnocování - tvorba pylových diagramů
Soubor pylů a spor v jednom vzorku, které pod mikroskopem determinujeme a
kvantitativně vyhodnocujeme, se označuje jako pylové spektrum.
Pylová spektra z chronologicky odebraných vzorků se znázorní v pylovém diagramu.
Získáme tak obraz časových změn vegetace na studovaném místě. Nejčastější
vyhodnocování pylových spekter se provádí na základě procentuálního zastoupení
jednotlivých palynomorf nebo jejich skupin. Nejmenší suma palynomorf v jednom vzorku,
ze kterých se pylový diagram vytváří, je 100 determinovaných zrn při minimálním množství
15 determinovaných taxonů. Menší množství není pro interpretaci vegetace dostatečně
reprezentativní.
Pro kvantitativní (semikvantitativní) vyhodnocování pylových spekter a tvorbu
pylových diagramů existuje několik softwarových programů. Jsou založené na
procentuálním zastoupení determinovaných taxonů v jednotlivých vzorcích. V mé práci
21
bylo využíváno především programu POLPAL polských autorů Walanus, Nalepka (1999).
Tento program umožňuje jednoduše a operativně vložit do tvorby pylových diagramů i
zkušenosti palynologa. Tohoto využívám např. při kombinování taxonů s ekologicky
podobnými nároky do jedné křivky tzv. syntetického diagramu, což v mnoha případech
umožní názornější vizualizaci ekologických charakteristik prostředí (např. Doláková et al.
2010, 2014). Toto kombinování má ovšem svá omezení, protože ne vždy se podaří dané
taxony determinovat až do úrovně druhu, což může způsobit nepřesnosti u těch taxonů, které
mají různé ekologické nároky pro různé druhy (např. Quercus – dub – sušší stanoviště nebo
lužní lesy, existence stálezelených druhů v tercierních palynospektrech). Ještě složitější je to
u taxonů tercierních, kde u některých morfotaxonů není stoprocentně objasněná botanická
příslušnost, a tudíž ekologická charakteristika.
Pylové taxony, které jsou v asociaci výrazně nadhodnocené (akumulují se v důsledku
vysoké pylové produkce a tafonomických procesů), a tudíž by výrazně změnily poměr
jednotlivých rostlin, je možné vyloučit z celkové sumy 100%. Graf je potom možné sestavit
ze dvou částí. Levá část znázorňuje procentuální zastoupení jednotlivých taxonů nebo jejich
skupin, kde se za celkovou sumu počítá součet všech pylových zrn s vyloučením
nadhodnoceného taxonu. Pravá strana grafu znázorňuje poměr zastoupení původně
vyloučeného taxonu k celkové sumě všech determinovaných zrn. Botanikové používají tento
princip pro odstranění vlivu ekologicky výrazného stanoviště nad celkovým charakterem
širšího okolí (např. převaha Cyperaceae, Polypodiaceae). V mé praxi se tento princip
osvědčil pro interpretaci pylových spekter zejména jeskynních nebo mořských sedimentů,
kde dochází k selektivním změnám v důsledku tafonomických procesů (Doláková 2014,
Doláková et al. 2014).
3. Palynologie neogénu
Studium palynologie v sedimentech neogenního stáří (mladší terciér) tvoří největší
část mé palynologické práce. Publikované výsledky se soustřeďují na 2 vzájemně propojená
témata:
3a) rekonstrukci vegetačních poměrů v jednotlivých časových úsecích, a na jejich základě
interpretace paleoekologických charakteristik a stratigrafických údajů;
3b) interpretaci širších geologických charakteristik na základě koordinace výsledků
dalších paleontologických, sedimentologických a geochemických metod. Tyto práce se
snaží interpretovat poměry v sedimentačních pánvích a řešit vzájemné vztahy
kontinentálního a mořského prostředí.
22
Studované lokality patří k velmi geologicky velmi dynamickému území, které je
situováno na hranici mezi východními svahy tektonicky klidné Evropské platformy
(reprezentované českým masívem) a okrajem tektonicky aktivních, zvyšujících se horských
řetězců Západních Karpat. Během neogénu bylo toto teritorium pokryto širokým výběžkem
epikontinentální mořské oblasti Centrální Paratethydy. Území mělo během studovaného
období velmi komplikovanou geodynamickou historii (Kováč 2000, Kováčová et al. 2011).
Vývoj morfologie krajiny se zásadním způsobem projevoval i ve změnách vegetace
zobrazené v pylových spektrech. Díky velké prostorové distribuci palynomorf lze pylová
spektra studovat i v mořských sedimetech a využít je ke korelaci kontinentálního a mořského
prostředí.
Publikované výsledky byly soustředěny na lokality geodynamicky odlišných lokálních
pánví Centrální Paratethydy: karpatské předhlubně, vídeňské pánve a jihoslovenské a
modrokameňské pánve na Slovensku.
Studovaný časový interval zahrnuje stupně standardní chronostratigrafické stupnice
burdigal až torton, které jsou na území Centrální Paratethydy označovány jako eggenburg
až pannon (obr. 6). Stratigrafické členění a hlavní typy sedimentů v karpatské předhlubni na
Moravě jsou uvedeny na obr.3. Sedimenty, ve kterých byla palynospektra studována,
zahrnují prostředí mořská, nebo různé okrajové vývoje mořské pánve s brakickou až
sladkovodní sedimentací, případně delty velkých řek i sedimenty tufitů. Výhodou studia
palynospekter v mořských sedimentech je jejich přesná chronostratigrafická datace na
základě mořských mikroorganizmů (foraminifera, Ca-nannoplankton). Toto datování potom
umožňuje korelace s klimatickými změnami pozorovatelnými v terestrických vývojích.
23
Obr. 6. Chronostratigrafie a biostratigrafie miocénu (Harzhauser, Piller 2007)
24
Obr. 7. Regionální stratigrafické členění neogénu karpatské předhlubně na Moravě (Rasser
et al. 2008)
Období mladšího terciéru – neogén je charakteristické řadou globálních i lokálních
klimatických změn. Změny globálního charakteru jsou zaznamenány především na
izotopových hodnotách kyslíku ze schránek organizmů z hlubokomořských vývojů. Přechod
z paleogénu do neogénu (oligocén- miocén) je vyznačený krátkou chladnou oscilací známou
jako event Mi-1, který byl zaznamenaný před 23,7 mil let (Zachos et al. 2001). Po tomto
ochlazení po celou dobu spodního miocénu teploty opět vzrostly. Menší klimatické oscilace
odpovídají vrcholům hodnoty izotopů kyslíku (Mi - eventy), které byly zaznamenány
opakovaně (Miller et al. 1991, Shackleton et al. 1999). Další teplotní maximum je známé
jako Mid-Miocene Climatic Optimum (MMCO), datované do období před 17 až 14,5
miliony let kolem hranice spodního a středního miocénu (Zachos et al. 2001). Toto období
představovalo nejteplejší časový interval za posledních 35 mil. let. Po něm následovala
25
celosvětově zaznamenaná fáze přechodného klimatu známá jako Middle Miocene Climatic
Transition (MMCT), které se projevovalo v rámci převládajícího globálního klimatu před 16
a 14.8 mil. let jako krátkodobé oscilace klimatu, objemu Východoantarktického
ledovcového štítu (EAIS), hladiny světového oceánu a hlubokooceánské cirkulace.
V pozdější fázi, od 14,8 - 12,9 mil. let, klimatický vývoj zahrnoval velký vzrůst EAIS
spojený s ochlazením Antarktidy, znatelný růst teplotního gradientu, rozsáhlé výkyvy
mořské hladiny následované globálním snížením hladiny oceánu a významné změny
hlubokomořské cirkulace (Flower and Kennett 1994). V mělkomořských okrajových
vývojích byl klimatický záznam kromě globálních podmínek ovlivněn i lokálními
geomorfologickými podmínkami a vulkanickou aktivitou během vývoje Alpskokarpatského
horského pásma a rovněž průniky chladných nebo teplých oceánských vod
(Holcová et al. 2015, Kováč et al. 2017).
Terestrická vegetace tyto změny odráží velmi výrazně. Stratigrafie kontinentálních
vývojů (kde chybí indexové fosílie) střední Evropy je proto do značné míry založena na
studiu těchto klimatických změn (klimatostratigrafie).
3a) Interpretace vegetace a klimatických změn
V palynospektrech z území ČR bylo v období neogénu zachyceno mnoho
klimatických výkyvů, které jsou typické pro centrální Evropu. Jejich interpretací
v terestrických ekosystémech a stratigrafickým zařazením se zabývala a zabývá celá řada
autorů (např. Mai 1981, 1991, Planderová 1990, Planderová et al. 1993a,b, Sadowska 1989,
1993, Stuchlik 1992, Stuchlik et Ważynska 1993, Mosbruggeret Utescher 1997, Böhme
2003, Böhme et al. 2006, Bruch et al. 2004, 2007, Kvaček et al. 2006, Kováčová et al.
2011)
Celkový charakter klimatu, zejména kombinace teplotních a srážkových poměrů, je
zobrazen v tzv. zonální vegetaci (odráží hlavní klimatickou zonaci – tropické deštné lesy,
opadavé lesy mírného pásma apod.). Obraz místních paleoekologických a
paleogeografických poměrů, často závislých na charakteru substrátu, vyjadřují společenstva
azonální (intrazonální x extrazonální, např. vegetace lužních lesů, slaniska, horská vegetace).
Azonální vegetační prvky mohou být v palynospektrech lokálně značně nabohacené a
mohou zastírat celkový zonální charakter. Lze z nich ovšem interpretovat lokální poměry
daného regionu a rovněž vztahy mezi mořským a kontinentálním prostředím (zasolování
pobřeží, ingrese sladkých vod, mírný nebo diferencovaný reliéf pobřeží - vznik pobřežních
26
bažin, horská vegetace). Odlišit je navíc třeba, pokud možno i taxony nabohacené
v důsledku rozdílné transportní schopnosti palynomorf a tafonomických vlivů (viz kap.
2a,b). Další problém v interpretaci vegetačního pokryvu a klimatických faktorů představují
morfotaxony, u nichž je dosud botanická příslušnost systematicky nejistá. Pro interpretace
palynospekter proto není možné bezvýhradně využít aktualistických údajů. Pro přesnější
představy o paleoprostředí jsou nutné poznatky dalších paleontologických,
sedimentologických a geochemických metod.
Změny klimatických i dalších paleekologických charakteristik se projevují nejvíce ve
vzájemném poměru zastoupení méně teplotně náročných rostlin (reprezentovaných zejména
opadavými dřevinami a jehličnany) – tzv. arktotercierních geofloristických prvků (A) a
rostlin s vyššími klimatickými nároky (stálezelené velkolisté taxony rostoucí v podmínkách
tropického a subtropického pásma) – tzv. flóry paleotropických geofloristických prvků (P).
Tento princip jako první navrhl Mai (1981, 1991). Palynologická společenstva
charakterizující jednotlivé klimatické změny označil jako pylové obrazy. Stuchlik et al.
(1994) rozdělili arktoterciérní taxony (A) na teplé mírné (A1) a chladné mírné (A2).
Paleotropické elementy (P) dělí na prvky klimatické zóny tropické (P1) a subtropické (P2).
U taxonů, které dosud nemají determinovanou botanickou příbuznost, posuzují ekologickou
náročnost na základě celkové charakteristiky společenstev, v nichž se tyto taxony vyskytují
(Stuchlik et al. 1993, 1994, 2014).
První komplexní interpretace a charakterizace klimatických změn na základě
vegetačních poměrů pobřeží Centrální Paratethydy byla shrnuta v pracích pod editorstvím
E. Planderové (Planderová et al. 1993a,b). K těmto výsledkům přispěly práce mnohých
autorů z jednotlivých území např. Konzalová (1976), Konzalová, Stuchlik (1983, 1992),
Nagy (1969, 1985), Planderová (1971, 1984 1990), Sadowská (1989, 1993), Zdražílková
(1993) a další. Autoři vymezili v miocenu Centrální Paratethydy čtyři výrazná
paleoklimatická období.
První interval zahrnující svrchní eger, eggenburg a ottnang je charakterizovaný
tropicko – subtropickým klimatem se stále přetrvávající flórou paleogenního charakteru, s
typickými paleogenními taxony Cicatricosisporites chattensis, Plicatopollis plicatus,
vyskytujícími se jen v jižních oblastech Centrální Paratethydy. V severních oblastech byly
nalezeny paleotropické prvky ze skupiny P1 (Dicolpopollis kockeli, Fususpollenites fusus,
Sapotaceoidaepollenites), ale bez paleogenních elementů. Kromě zmíněných rozdílů se
pylová společenstva na celém území příliš nelišila. Tato shoda byla nejspíše způsobena
paleogeografickou situací území.
27
Druhý interval zahrnuje karpat a spodní baden. Jsou pro něj typické velmi hojné
paleotropické prvky a vyšší taxonomická diferenciace. Početné jsou taxony P1 i P2. Spodní
baden znamenal pro některé charakteristické tropické elementy jejich poslední výskyt
(Sapotaceae, Symplocaceae, Schizacaceae).
Třetí interval charakterizuje svrchní baden a sarmat. Paleoflóra se stává jednotnou,
docházelo k promíchávání flóry nížin s flórou Paratethydy, což mělo za následek rozšíření
arktoterciérních prvků na jihu a vzrůstem paleotropických elementů skupiny P2 na severu.
V sedimentech z období svrchního sarmatu severně od Karpat převládá močálová
teplomilná vegetace s příznivými podmínkami pro tvorbu uhlí. Míšení flór mizí během
postupné regrese moře, paleotropická vegetace migruje na jih. Od tohoto období dominují
severně od Karpat prvky arktoterciérní.
Čtvrtý interval představují stupně pannon a pont. V tomto časovém úseku docházelo
k expanzi arktoterciérní flóry a dále ke vzrůstu kvantity chladnomilných taxonů v severní
části Paratethydy. Klimatické podmínky v pontu jsou mírné humidní a vegetace se vyznačuje
převážně listnatými lesy rozprostírajícími se u břehů teplých vod (Planderová et al. 1993).
Na tyto práce navazovaly některé další komplexní publikace, které problematiku
rozšiřovaly a upřesňovaly (např. Stuchlik et al. 1994, Nagy 1999).
V současnosti jednou z nejvyužívanějších metod pro interpretace paleoklimatu na
základě terestrických rostlinných společenstev je tzv. „koexistenční přístup“ - „coexistence
approach (CA)“ - který poprvé publikovali Mosbrugger, Utescher (1997). Rozsáhlou diskusi
jeho aplikací a problematiku využití zpracovali Utescher et al. (2014). Tato metoda je
založená na aktualistickém předpokladu, že tercierní taxony rostly v podobných podmínkách
jako jejich nejbližší žijící příbuzní (NLR – „nearest living relatives“). U těchto recentních
zástupců jsou definované hlavní ekologické faktory (ekologická valence), které jsou shrnuty
do databáze CLIMBOT (zahrnuje více než 800 tercierních rostlinných taxonů v podobě
výtrusů, pylových zrn, semen, listů a dřev, jejich nejbližší recentní příbuzné a jejich
klimatické parametry). Klimatická charakteristika jednotlivých taxonů vyjadřuje 10
hlavních parametrů - průměrná roční teplota (MAT), průměrná teplota v nejteplejším měsíci
(WMT), průměrná měsíční teplota v nejchladnějším měsíci (CMT), průměrné roční srážky
(MAP), minimální průměrné roční srážky (MinAP), maximální měsíční srážky (MmaP),
minimální měsíční srážky (MmiP), srážky v nejteplejším měsíci (WMP), relativní vlhkost
(RH), index aridity (AI). Výsledkem tohoto modelu je získání intervalových hodnot u
jednotlivých parametrů (teplotních, srážkových), v rozmezí kterých mohou determinované
taxony na určitém místě růst ve společné asociaci (koexistovat). Pro získání koexistenčních
28
intervalů byl navržen program CLIMSTAT (Mosbrugger, Utescher 1997). Jedny z prvních
konkrétních výsledků interpretací klimatických charakteristik a jejich vývoje během
posledních 25 mil. let vývoje klimatu na základě CA z oblasti SZ Německa publikovali
Utescher et al. (2000).
Databáze Palaeoflora (http://www.palaeoflora.de/) je neustále rozšiřována a je
součástí mezinárodního programu NECLIME - Neogene Climate Evolution in Eurasia, který
byl založen v roce 2000 (http://www.neclime.de/). Výsledky této rozsáhlé spolupráce
odborníků na neogénní terestrické ekosystémy jsou neustále publikovány a rozšiřovány
(např. Bruch et al. 2007, Utescher et al. 2011, Louis et al. 2017).
Jednou z prvních syntetizujících prací v rámci tohoto programu byla publikace Bruch
et al. 2004. Výsledkem CA analýz z mnoha lokalit střední a jižní Evropy z období časného
středního miocénu a časného svrchního miocénu byla rekonstrukce klimatického gradientu.
V obou těchto obdobích byl konstatován pouze mírný šířkový gradient. Tento jev patrně
souvisel s obrovským plošným rozsahem epikontinentální mořské oblasti Centrální
Paratethydy, která klimaticky ovlivňovala suchozemské oblasti ve svém okolí. Detailnější
klimatické interpretace související např. s rozdílnou paleogeografií v důsledku zvyšujících
se oblastí alpsko karpatského horského pásma jsou patrně studovanou metodou
nezachytitelné.
Z prostředí karpatské předhlubně byla do klimatických modelů zahrnuta lokalita
Židlochovice, kde byly na několika palynospektrech z bodových vzorků vypracovány
základní klimatické charakteristiky. Tato lokalita reprezentuje hypostratotyp spodního
badenu, a byla proto zpracována podrobněji ve dvou 20 metrových vrtech
z paleontologického i sedimentologického hlediska a rovněž metodou CA (Doláková et al.
2014) viz níže.
Jako každá interpretační metoda má i CA ve fosilním záznamu svá omezení. Je
založena pouze na přítomnosti nebo nepřítomnosti určitého taxonu v paleoasociaci.
Nezohledňuje kvantitativní zastoupení některých prvků a vlivy tafonomie – nedochování se
taxonů s rychleji rozložitelnou stěnou, nebo taxonů, které se vlivem menší schopnosti
distribuce palynomorf nedostanou do studovaného sedimentu, např. mořského prostředí. U
tercierních palynospekter, kde je vždy určitý podíl pouze morfologicky determinovaných
taxonů, je navíc interval koexistence značně široký. Chyby v interpretaci mohou být u
tercierních taxonů rovněž způsobeny nepřesným přiřazením nejbližšího příbuzného,
případně změnami ekologické valence některých taxonů - např. určitý druh se vyskytoval
v odlišných asociacích než dnes – refugia. Příkladem může být rod Craigia. Tento rod je v
29
současné době zastoupený v asociaci širolistých stálezelených smíšených lesů blízkých
tropům, se sezónními zvýšenými srážkami z Číny a severního Vietnamu. V miocenních
oryktocenózách se však ve vyšších frekvencích vyskytoval v asociacích pobřežních až
uhlotvorných bažin v asociaci opadavých dřevin (s převažujícími Taxodioideae). (Kvaček et
al. 2002, 2004, Zetter et al. 2002). V těchto asociacích se pylová zrna rodu Craigia
vyskytovala hojně i ve spodním miocénu karpatské předhlubně na Moravě (Doláková 2004,
Kováčová et al. 2011)
Práci, která propojuje poznatky vegetačních poměrů a geodynamického
vývoje z prostoru Centrální Paratethydy, představuje publikace Kvaček et al. (2006). Pro
interpretaci a zobrazení vegetačních a paleogeografických poměrů bylo využito digitálního
elevačního modelu (DEM). Tento model umožňuje zobrazit představu krajiny na
palinspastických mapách s rekonstruovanou orografií vývoje karpatského horstva a okolních
sedimentačních pánví.
Model vegetace kontinentálního prostředí v okolí Centrální Paratethydy využívá
zjednodušený systém vegetačních jednotek (formací), které jsou postačující pro získání
předběžného obrazu paleovegetace v prostoru.
V rámci mezinárodního týmu byly (na základě studia listové flóry, semen, plodů a
palynomorf) pro rekonstrukci rostlinného pokryvu navrženy základní zonální, intrazonální
a extrazonální formace (vegetační jednotky). Tyto formace sdružují jednotlivé elementy
flóry na základě autekologie jejich nejbližších žijících příbuzných (NLR) a rovněž metod
listové fyziognomie. Rostlinná společenstva referenčních fosilních lokalit často obsahovala
taxony rozdílných vegetačních formací, z nichž zonální prvky byly použity pro tvorbu map
rekonstrukcí fosilní vegetace.
Charakteristika formací byla založena na diverzitě a vzájemném poměru zonálních
elementů: stálezelených, opadavých, sklerofylních a leguminozních dřevin. Kromě zonální
vegetace byly shrnuty i základní typy vegetace azonální (intrazonální, extrazonální).
Zonální formace – procentuální zastoupení je poměr ke všem determinovaným
zonálním krytosemenným.
1. Temperátní listnatý (širolistý) opadavý les - (Warm-) temperate Broad-leaved
Deciduous Forest) s velmi nízkým poměrem stálezelených dřevin, obsahující více než 80%
opadavých zonálních krytosemenných dřevin (Parrotia, Zelkova, Ostrya, Acer
angustilobum atd).
2. Temperátní smíšený mezofytní les (Warm) - temperate Mixed-Mesophytic Forest zahrnující
méně než 80% zonálních opadavých krytosemenných dřevin, méně než 30%
30
stálezelených krytosemenných dřevin, méně než 20% sklerofylních a leguminozních
(malolistých) typů s pravidelnou příměsí Tetraclinis salicornioides a dalších teplomilných
prvků, méně než 30% zonálních bylin)
3. Subtropický širolistý stálezelený les - Subtropical Broad-leaved Evergreen Forests,
zahrnující tzv. mastixiovou flóru “Younger Mastixioid Floras” sensu Mai (1981), obsahující
30 % nebo více listnatých stálezelených teplomilných elementů, reprezentovaných zejména
čeleděmi Lauraceae, Theaceae, Mastixiaceae, Symplocaceae, Sapotaceae, Engelhardia, a
stálezelené členy čeledi Fagaceae (v pylovém spektru reprezentované morfologickými
druhy Castaneoideoipollenites pusillus, Quercoidites henrici, Quercoidites microhenrici) a
méně než 25% zonálních bylin.
4. Subtropický a subhumidní sklerofytní les - Subtropical sub-humid Sclerophyllous
Forest, obsahující více než 20 % sklerofylních taxonů (Quercus mediterranea, Quercus
drymeja) a leguminozních malolistých dřevin
Azonální formace – význačné množství těchto prvků ve společenstvech může zastřít
celkový charakter klimatických zón - tzv. faciálně ovlivněná společenstva, nejsou vyjádřená
v mapách pomocí vzorců, ale pouze bodově u referenčních lokalit.
Intrazonální formace:
5. Bažinný les a uhlotvorný močál - Swamp forest and coal-forming mire, kde
dominantní postavení mají elementy uhlotvorných dřevin a bylin (Glyptostrobus adalších
Taxodioideae, Byttneriophyllum, Nyssa, Myrica, Calamus, Spirematospermum).
6. Močálová a akvatická vegetace - Marsh and aquatic vegetation, zastoupená
dominantně vodními bylinami a halofyty (Cyperaceae, Typha, Potamogeton, Stratiotes atd.).
7. Opadavý lužní les - Deciduous riparian forest, tvořený především dřevinami
vlhkých substrátů (Taxodium, Alnus, Salix, Populus, Fraxinus, Acer tricuspidatum)
Extrazonální formace
8. Horské jehličnaté lesy - Mountain conifer-rich forest reprezentované především
rody Cedrus, Tsuga, Picea, Cathaya etc. Tato formace je v mapách znázorněná, je založená
většinou na záznamu pylových zrn.
Na základě těchto typů vegetace byly výsledky floristických studií analyzovány ve 3
časových intervalech miocénu s odlišnou flórou a paleogeografií a byl z nich vytvořen
paleobiogeografický model pro areál Centrální Paratethydy a její periferii. Jednalo se o
inervaly : karpat/spodní baden, svrchní baden/spodní sarmat a spodní panon.
Limity modelu spočívají v tom, že nebere v úvahu hojnost výskytů, což bylo
způsobeno nepravidelným fosilním záznamem. Nicméně procentuálně vyjádřená diverzita
31
je prozatím nejobjektivnější charakteristika rostlinných společenstev. Na
paleogeografických podkladech úzkých časových úseků lépe vynikne dynamika vegetace a
její vývoj, i když je pro daný čas malý počet referenčních lokalit. Problematická byla
transformace palynologických diagramů, ve kterých zůstává stále určitý počet taxonů
s pouze morfologickými determinacemi do jednotlivých vegetačních formací, které byly
vytvořeny na základě zejména listové fyziognomie. Pokud to bylo možné, byly starší
palynologické údaje revidovány, aby se rozšířilo množství taxonů přiřazených do
botanického systému. Mnohá zpřesnění ovšem zůstávají otázkou pro budoucí výzkumy.
3ab) Charakteristika vegetace v jednotlivých časových úsecích
Prozatím nejnovější kompletaci a interpretaci palynologických údajů doplněnou o
literární údaje makrofloristické z území ČR a SR a srovnání s vývojem flóry v sousedních
oblastech představuje pulikace Kováčová, Doláková a Kováč (2011). Práce se zaměřila na
vegetační a paleoklimatickou analýzu dvou oblastí Centrální Paratethydy a jejího okolí
s různým geodynamickým vývojem: karpatské předhlubně na Moravě a vídeňské pánve na
Moravě a na Slovensku. Shrnuje výsledky dosavadních výzkumů zaměřených na vegetaci
miocénu studovaných areálů (Doláková 2004, Doláková et al. 1999, Doláková, Kováčová
2008, Doláková et al. 2011, Doláková, Slamková 2003). Výsledky pylové analýzy
výcházejí ze semikvantitativního hodnocení palynospekter na základě proporcionálního
poměru paleotropických a arktotercierních prvků (sensu Mai 1981, 1991, Stuchlik et al.
1994, Neclime koncept NLR.), které jsou vyjadřovány pomocí pylových diagramů.
Reprezentují vývoj vegetace a klimatu ve sledovaných stratigrafických intervalech.
Interpretace výsledků vychází z konceptu prací Kvaček et al. (2006), Kovar-Eder et al.
(2008a, b).
Na změnách pylových spekter se kromě klimatického vývoje projevují i změny
paleogeografické, jako např. rozšiřování a restrikce mořských pánví – transgrese a regrese
spojené se zarovnáváním reliéfu nebo silnou erozí kontinentu, vznik a vzájemné propojování
lokálních okrajových pánví, prostředí delt s míšením mořské a sladké vody a přínosem
organické hmoty z pevniny, zvyšování reliéfu v důsledku orogeneze, dosouvání karpatských
příkrovů – nedochování okrajových facií apod. (Kováč 2000, Nehyba et al. 2008).
Z konceptu práce Kováčová, Doláková, Kováč (2011) vyplynuly peleovegetační
chjarakteristiky stratigrafických úseků miocénu a byly doplněné údaji z některých starších i
novějších vlastních publikací.
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Základní taxony, které jsou zahrnuty do zonální vegetace skupiny teplomilných
stálezelených dřevin (P), jsou - Sapotaceae, Palmae, Engelhardia, Platycarya, stálezelené
členy čeledi Fagaceae (morfotaxony Quercoidites microhenrici a Quercoidites henrici,
Trigonobalanopsis), Symplocos, Cornaceae - Mastixiaceae, Tricolpopollenites liblarensis,
Araliaceae, Rutaceae, Reevesia, Alangium, Theaceae, Styracaceae, Parthenocissus, atd.
Základní prvky opadavých dřevin – arktotercierní (A) vegetace jsou: Quercus, Acer,
Carpinus, Betula, Juglans, Tilia, Celtis, Carya, Zelkova, Ostrya, Eucommia.
Extrazonální horská vegetace je reprezentována rody: Cedrus, Tsuga, Picea, Cathaya,
Keteleeria.
Azonální vegetace ovlivňovaná edafickými faktory je reprezentována:
1. prvky lužního lesa: Alnus, Salix, Ulmus, Fraxinus, Pterocarya, Platanus,
Liquidambar, Cercidiphyllum;
2. pobřežních bažin: Taxodioideae (Glyptostrobus), Myricaceae, Cyrillaceae, Nyssa,
Craigia, Sciadopitys, Decodon, Ilex;
3. vodní a příbřežní vegetace: Cyperaceae, Oenotheraceae, Sparganium, Typha,
Potamogeton, Nelumbo, Azolla;
4. vegetací otevřených ploch se zástupci heliofyt: stromovitých Olea, bylinných
(Poaceae, Caryophyllaceae, Asteraceae, Artemisia, Salvia) a keřovitých (Ericaeae, Buxus,
Rosaceae, Ephedra) a pobřežních slanisek (Chenopodiaceae – Salicornia and Gypsophilla,
Tamarix)
5. kapraďorosty – teplomilné: Lygodium, Pteris, Schizeaceae, Davaliaceae,
ostatní: Polypodiaceae, Osmunda, Selaginella.
Prostředí spodnomiocenních stupňů eggenburg – ottnang (obr. 6,7) zachycené
v sedimentech jz. části karpatské předhlubně bylo mimořádně variabilní (např. vrty
Miroslav, Jezeřany, Šafov, Čejkovice). Sedimentace probíhala v subtropickém klimatu
v příbřežní mořské oblasti s velmi morfologicky diferencovaným reliéfem pobřeží – mořské
facie se střídaly v čase i prostoru s faciemi lagun a delt (brakické až sladkovodní prostředí).
Změny mořského prostředí byly spojené se změnami mořské hladiny, změnami dynamiky,
prosvětlení, obsahu kyslíku a salinity a rovněž rychlostí akumulace sedimentů. Tyto změny
se projevily jak v mořských ekosystémech, tak i v charakteru pobřežní terestrické vegetace
(Doláková et al. 1999, Nehyba et al. 1995, 1997, Doláková 2004). Palynospektra měla
charakter subtropického klimatu. V porovnání s mladšími obdobími miocénu jižní části
karpatské předhlubně byla typická pravidelná přítomnost pylových zrn čeledi Rutaceae,
33
hojněji byl zastoupený subtropický rod Symplocos a ojediněle byla zjištěna pylová zrna rodu
Alangium. Ve facii lužních lesů byl zastoupený hojněji rod Platanus než Alnus. Vegetace
otevřených ploch se zástupci heliofyt (Poaceae, Caryophyllaceae, Asteraceae, Ericaeae,
Buxus, Ephedra) a pobřežních slanisek (Chenopodiaceae až 37% ve vzorku, patrně rod
Salicornia, Tamarix) se v profilech střídala s převahou rostlin pobřežních bažin
(Taxodioideae, Myricaceae, Cyrillaceae, Decodon) s hojným podílem teplomilných
kapradin. Např. rod Lygodium (až 5 %), Pteridaceae, Gleicheniaceae, vranečky Selaginella
a jatrovkou Riccia. Flóra rostoucí na okrajích sladkých vod (Sparganium, Potamogeton,
Nelumbo, Cyperaceae) byla rovněž charakteristická a častěji zastoupená než ve spektrech
karpatu a badenu. Typická pro palynospektra eggenburgu byla přítomnost formálního rodu
Monocirculipollis, který nebyl zjištěn v mladších sedimentech karpatské předhlubně. Na
základě studia v elektronovém mikroskopu byl přiřazen čeledi Caryophyllaceae (zmíněno
v kapitole 2cb), morfologicky podobnému pylovým zrnům rodu Gypsophilla. Vysoká
frekvence jehličin čeledi Pinaceae zahrnující nejčastěji rody Pinus, Cathaya a méně Cedrus,
Picea, Abies prokazovala velmi diferenciovaný reliéf pořeží. Rod Tsuga byl na rozdíl od
mladších sedimentů zjištěn pouze velmi ojediněle.
Nálezy palynologie podpořily dřívější závěry studia makroflóry, která je ve studované
oblasti poměrně vzácná. Ojedinělou oryktocenózu ze Znojma a Přímětic determinoval
Knobloch (1982, 1984). Interpretoval několik ekologicky odlišných asociací: a) keřovitobylinnou
heliofilní vegetaci se zástupci stálezelených sklerofylních dřevin podobnou
mediteranním společenstvům tzv. macchie; b) bažinnou vegetaci Glyptostrobus, Myrica; c)
akvatickou flóru se Salvinia, Potamogeton, Nymphaea a pobřežní společenstvo s Typha,
Decodon, Sparganium. Popsal rovněž akumulaci plodů rodu Limnocarpus, který roste na
okrajích brakických vod.
Interpretace charakteru vegetace obsažené v sedimentech karpatu byla zpracovávána
zejména v rámci mezinárodního projektu Reedice stratotypu Karpatu. Závěry tohoto
projektu byly shrnuty v monografii Brzobohatý et al. (2003). Palynospektra byla mimo
dalších vrtů a lokalit studována i na stratotypových lokalitách Slup, Hevlín a Dolní
Dunajovice (Doláková, Slamková 2003, Doláková et al. 2003).
Podle palynologických údajů i dalších geologicko-paleontologických studií
(Brzobohatý et al. 2003, Kvaček et al. 2006) byla pobřežní oblast Centrální Paratetydy
charakteristická nízkým zarovnaným reliéfem, který představoval příznivé podmínky pro
tvorbu hojných bažin v příbřežním a deltovém režimu. Během karpatu nebyla na základě
34
paleobotanického studia patrná výrazná šířková ani výšková vegetační zonace v celé karpato
- panonské oblasti
Základní charakter zonálních společenstev vyjádřený v poměru teplomilných
paleotropických prvků vůči prvkům arktotercierním se oproti eggenburgu a ottnangu
výrazně nezměnil. Převažující formaci tvoří subtropické listnaté lesy s vysokým nebo
středním podílem stálezelených dřevin. Některé taxony jako Symplocos, Rutaceae a
Platanus byly oproti předchozímu období nalézány sporadičtěji. Oryktocenózy bažinných
facií (Myricaceae, Taxodioideae, Craigia, Pteridaceae, Polypodiaceae) a lužních porostů
(Alnus, Ulmaceae, Lythraceae) se vyskytovaly běžně a často tvořily převažující složku
palynospekter. Velmi typické byly asociace s výrazným zastoupením rodu Craigia.
Obdobné oryktocenózy byly pozorovány v cyprisových jílovcích ze Severočeské pánve pod
formálním taxonem Intratriporopollenites insculptus Mai (Konzalová (1976) a v
karpatských sedimentech lokality Korneuburg (Hofmann et al. 2002).
Zástupci bylinné a keřovité vegetace otevřených stanovišť byli daleko méně
zastoupeni než v období předcházejícím. Typické mořské sedimenty - tzv. “šlír” - byly
charakteristické vysokým podílem jehličin, zejména rody Pinus a Cathaya, které se díky
velkému doletu pylových zrn těchto větrosnubných rostlin často hromadí v mořských
sedimentech (viz kap. 2). Rody Tsuga a Abies jako zástupci extrazonální vegetace rostoucí
převážně ve výše položených oblastech se začínají objevovat v sedimetech z mladších
období studovaného intervalu. Dokumentují počátek výzdvihu karpatského orogénu
(Doláková et al. 2003, Kvaček et al. 2006, Kováčová et al. 2011).
Vyskytovali se rovněž zástupci mořských dinofyt a cysty zelených řas skupiny
Prasinophyta, které byly místy nalézány v masovém množství. Tyto akumulace mohly být
projevem vodního květu, který se vytváří v důsledku omezené cirkulace a stratifikace
vodního sloupce. Pylová zrna a spory byly často degradovány a obsahovaly shluky
krychlových dutin (pozůstatky po mikrokrystalech pyritu), které svědčí o existenci prostředí
se sníženým obsahem kyslíku (viz kap. 2).
Existence subtropického humidního klimatu byla podpořena i makrofloristickými
nálezy na stratotypových lokalitách Slup a Dolní Dunajovice. Asociace byly tvořeny
převahou fosilních listů čeledi Lauraceae (pylová zrna této čeledi se nedochovávají ve
fosilním stavu) a nízkým zastoupením opadavé flóry (Knobloch 1967, 1982, Kvaček 2003).
Azonální vegetace byla reprezentována bažinnými a lužními lesy s dominancí rodů
Glyptostrobus a Myrica. Zjištěny byly i další prvky totožné s nálezy pylových zrn včetně
rodu Craigia.
35
Sedimenty spodního miocénu ottnang - karpat byly z palynologického hlediska
studovány i z území Modrokameňské pánve na Slovensku (Doláková 2004). Charakter
oryktocenóz převážně odpovídal pylovým spektrům z karpatské předhlubně. Nejvýraznější
rozdíl představovalo zastoupení formálního rodu Pentapollenites Krutzsch, který v
palynospektrech z území Moravy nebyl zjištěn. Asociace palynomorf poněkud chladnějšího
klimatu reprezentovaného zvýšením poměru arktotercierních dřevin popsané z konce
ottnangu a počátku karpatu ze sousedních oblastí např. Slovenska, Polska (Planderová 1990,
Planderová et al.1993a,b, Ważyńska 1998), nebyly v karpatské předhlubni zaznamenány.
Tato palynospektra mají ovšem převahu azonálních prvků a změny mohly být odrazem
lokálních paleogeografických poměrů (vyšší diferenciace reliéfu).
Velmi obdobný charakter karpatských palynospekter byl publikován ze sedimentů
pohoří Mecsek z Maďarska (Nagy 1999). Palynospektra z karpatských sedimentů
Korneuburgské pánve v Rakousku vykazovala obdobný charakter zejména azonálních
formací. Vysoké poměrné zastoupení palem a nálezy rodu Avicenia, který představuje
ojedinělý prvek mangrovových porostů v oblasti Centrální Paratethydy, interpretují autoři
Hofmann et al. (2002) jako důsledek průniku teplých vod z oblasti Tethydy.
Podle nejnovějších studií a statistického vyhodnocení palynospekter pomocí CA
(Doláková et al in prep, jsou na pylových diagramech pozorovatelné cyklické změny
poměrného zastoupení teplomilných a arktotercierních prvků. Tato periodicita pokračuje i v
následujícím stupni baden.
Ve středním miocénu, který je na území karpatské předhlubně zastoupený stupněm
baden, byl vývoj krajiny ovlivněný zejména postupným výzdvihem horských oblastí
Západních Karpat a současnou subsidencí přilehlých nížinných oblastí. V palynospektrech
je patrný nástup výrazné výškové vegetační zonace (Kováčová et al. 2011).
Interpretace vegetace v sedimentech spodního badenu karpatské předhlubně byla
publikována zejména v pracích (Doláková 2004, Doláková et al. 2011, 2014, Kováčová et
al. 2011).
Palynologická data dokumentují subtropické klima v období doznívajícího
miocenního klimatického optima (MMCO) a dominancí zonální formace stálezelených
širolistých lesů (až přes 30% - Engelhardia, Platycarya, stálezelené typy Fagaceae:
Castanopsis, Trigonobalanopsis). Podíl prvků formace temperátního smíšeného
mezofytního lesa a širolistého opadavého lesa byl nižší (např. Quercus, Carya, Celtis,
Juglans). Oproti palynospektrům spodního miocénu byl zaznamenaný mírný úbytek
některých teplomilných prvků (P1 - Sapotaceae, Palmae, Mastixiaceae, Lygodiaceae).
36
V pylových spektrech byla pozorovaná vyšší diverzita pylových zrn stálezelených i
opadavých dubů a nárůst podílu pylových zrn rodu Platanus.
Kolísání podílu prvků lužního lesa a pobřežních močálových biotopů a místně zvýšený
podíl bylin a heliofytních rostlin vyskytujících se na sušších otevřených areálech byl
důsledkem změn humidity. Toto zjištění odpovídá i studiím makrofloristickým. V porovnání
s nálezy makroflóry (Knobloch et Kvaček, 1996) indikují palynologické údaje poněkud
teplomilnější charakter vegetace. Tento rozdíl může být ovšem způsoben tafonomickými
procesy v mořských sedimentech, kde se zejména listová flóra hůře dochovává. Zajímavý
fenomén představuje poměrně pravidelná přítomnost rodu Cercidiphyllum, který ve starších
sedimentech karpatské předhlubně nebyl dokumentován.
Občasné nálezy rostlin typických pro okraje sladkých vod, jako např. Potamogeton,
Sparganium a kolonie řas (Botryococcus), dokumentují přítok sladkých vod do mořské
pánve.
Největší pozornost byla věnována lokalitě Židlochovice. Jako na první lokalitě ze
studovaného prostoru byly statisticky spočítány na několika bodových vzorcích výsledky
CA (Bruch et al. 2004). Lokalita reprezentuje hypostratotyp sedimentů spodního badenu a
byla proto zpracována nověji z co nejširšího paleontologického (foraminifery, CA
nanoplankton, otolity, měkkýši, červené řasy, mechovky, palynomorfy) a
sedimentologického hlediska (Doláková et al. 2014). Problematika změn
paleobiologických, paleoklimatických a sedimentologických podmínek byla pro tuto
lokalitu zpracována kontinuálně ve dvou vrtech. Pylová spektra s bohatým a
diferencovaným obsahem palynomorf byla zpracována za pomoci CA. Ne všechny
studované vzorky byl ovšem k účelům statistického zhodnocení vhodné (viz kap. 2)
Na lokalitě Židlochovice byly ve dvou studovaných vrtech provedeny výpočty
teplotních a vlhkostních charakteristik za pomoci CA. Zvyšující se kolísání klimatických
charakteristik (zejména teploty a humidity, zvyšující se sezonalita) v rámci subtropické
kontinentální vegetace představují finální fázi vrcholícího miocenního klimatického optima
(MMCO) a nastupující středně miocenní přechodnou klimatickou fázi (MMCT) (Doláková
et al. 2014, Holcová et al 2015, 2018) ve smyslu Zachos et al. (2001), Flower , Kennet
(1994). Palynologická studia a studium asociací červených řas napomohla rovněž k
indikování prvního šířkového teplotního gradientu od spodního badenu v rámci pobřeží
Paratethydy (Doláková et al. 2008). Do tohoto období je předpokládána uniformita
charakteru klimatu v rámci celé střední Evropy (Bruch et al 2004).
37
Pro studium palynospekter svrchní části středního miocénu (svrchní baden, sarmat)
nebyly dosud v moravské části karpatské předhlubně (sedimentace končí ve spodním
badenu) ani vídeňské pánve nalezeny sedimenty vhodné k zachování pylových zrn a spor.
Všechny dosud zkoumané sedimenty byly palynologicky sterilní.
Podstatný nárůst taxonů horské vegetace (Picea, Abies, Tsuga, Cedrus), zvýšení
podílu arktotercierní flóry (Quercus, Ulmus, Carya) a úbytek subtropických taxonů
(Platycarya, Engelhardia, Myrica, Distylium, teplomilné Fagaceae, Sapotaceae)
v palynospektrech svrchního badenu byl dokumentován na lokalitách Slovenské části
vídeňské pánve. Tyto změny jsou interpretovány jako důsledek vzrůstu výškového gradientu
souvisejícího s výzdvihem horského oblouku Západních karpat (Kováč et al. 1998,
Doláková et al. 2011, Kováčová et al. 2011).
Studovaná palynospektra svrchního miocénu moravské části Paratethydy pocházela
ze sedimentů panonu vídeňské pánve. Vlivem paleogeografických změn se během svrchního
miocénu začalo postupně uzavírat spojení s mořskou oblastí Paratethydy. Severní část
vídeňské pánve byla postupně vyplňována progradujícími deltovými a říčními faciemi, které
přinášely materiál z vyzdvihujících se karpatských horstev. Prostředí se postupně
změlčovalo a vyslazovalo (Kováč et al. 1998a,b, 2017). Tento proces byl patrný i ze studia
palynologie, kde sporadické výskyty dinoflagelát a zelených řas čeledi Tasmanaceae indikují
ještě mořské až brakické prostředí. Místy hojné kolonie rodu Botryococcus se mohou
vyskytovat jak v brakické, tak sladké vodě. Ovšem kolonie rodu Pediastrum, cysty
Mougeotia, sporangia s glochidii vodní kpradiny Azolla, a pylová zrna pobřežních vodních
rostlin jako Nelumbo, Nymphaea, Myriophyllum, Sparganium, Potamogeton reprezentují
prostředí sladkých vod (Doláková, Kováčová 2008, Kováčová et al. 2011).
Sedimenty pannonu jsou známé bohatými makrofloristickými nálezy (Knobloch 1962,
1963, 1968, 1969, 1972, 1981, 1985), proto jim byla z hlediska palynologie věnována
pozornost již v dřívějších výzkumech (Gabrielová 1966, Kalvoda 1979, Lázničková 2006,
Konzalová 2005). V palynospektrech pannonských sedimentů byla oproti spodnímu a
střednímu miocénu zaznamenána výrazná změna vegetačního pokryvu. V důsledku
paleogeografických změn a klimatických oscilací se množství termofilních taxonů začalo
snižovat a některé z tohoto prostoru zcela ustoupily do jižnějších oblastí. Morfologicky
pestrý reliéf horských pásem v okolních oblastech vytvořil podmínky pro rozšíření
temperátních smíšených mezofytních lesů (Quercus, Celtis, Tilia, Carya, Zelkova, Ostrya,
Carpinus, Juglans) s minimálním přimíšením stálezelených subtropických taxonů
(Engelhardia, Castanea, stálezelené Fagaceae) a extrazonální horskou vegetaci s jehličnany
38
Cedrus, Tsuga, Picea (Doláková, Kováčová 2008, Doláková et al. 2010, Kováčová et al.
2011).
V tomto období se lokálně začínají objevovat areály otevřené krajiny se stepním
charakterem, tzv. „open woodland“ – „open grassland“, které se projevují zvýšeným
procentem výskytu bylinných prvků (10 až 14 % složení spekter) - Poaceae, Artemisia,
Chenopodiaceae, Asteraceae, Lamiaceae, Polygonum, Daucaceae, Caryophyllaceae,
Thalictrum, Rumex, Valeriana, Dipsacaceae, Galium, Ranunculus) a keřovitých forem
(Buxus, Ericaceae, Ephedra). Zjištěny byly rovněž lianovité formy Vitaceae, Lonicera,
Rosaceae typ Rubus) a halofyty (Chenopodiaceae) indikující existenci lokálně zasolených
substrátů.
Azonální vegetace se v jednotlivých sekvencích velmi rychle střídaly v čase i prostoru.
Pylová spektra tvoří různé facie intrazonálních společenstev, jako např. zasolené příbřežní
louky tzv. marše s Chenopodiaceae, Ericaceae, Poaceae, Tamarix) lužní lesy (Alnus, Salix,
Betula, Liquidambar, Myrica, Nyssa), pobřežní močály s Taxodioideae, Nyssa, Myrica
střídající se s asociacemi otevřených až stepních ploch (tzv. artemisiové stepi).
Na základě pylových společenstev byly za pomoci CA analýzy ze slovenské části
vídeňské pánve vypočítány klimatické faktory prostředí (Doláková, Kováčová 2008).
Obdobná palynospektra (redukované pobřežní močály a izolovaná sladkovodní jezera
obklopená stepními porosty (s dominantním pelyňkem Artemisia) s řídkými dřevinami
popsala ze slovenské části Podunajské pánve Planderová (1972, 1990). Ve srovnání
s palynospektry z Maďarska konstatovala na slovenském území chladnější a sušší klima
(Nagy 1985, 2005, Planderová 1990). Pylová společenstva bohatá na zastoupení bylin
popsali ze svrchního pannonu Štýrské pánve Hoffmann, Zetter (2005).
3b) Širší problematika neogénu v korelaci palynologie a dalších
paleontologických, sedimentologických a geochemických metod výzkumu
Výhodou studia palynologie v mořských sedimentech je možnost korelace
palynologických interpretací s paleoklimatickými a paleogeografickými údaji, které
vycházejí ze studia dalších paleontologických, sedimentologických, nebo geochemických
metod. Proto je poslední dobou práce širšího kolektivu autorů zaměřena na komplexní
zpracování studovaných lokalit a tvorbu modelů mořského a terestrického prostředí a
jejichvzájemného ovlivňování.
39
Prozatím nejpodrobnějším studiím byly podrobeny sedimenty spodního badenu
nannoplanktonové zóny MNN 5 (obr. 6.). Výsledky palynologického studia byly začleněny
do celkového pohledu na vývoj prostředí. Interpretace palynospekter i tafonomické procesy
se odrazily nejen ve změnách charakteru vegetace, klimatu a morfologie kontinentu, ale i
v interpretacích režimu a morfologie mořské pánve (změny obsahu kyslíku, proudění,
hloubky, salinity (Doláková et al. 2014, Holcová et al. 2015, Nehyba et al. 2016).
Výsledky palynologických studií přispěly i k interpretaci vzniku karbonátových těles
mořského původu, které samy jsou palynologicky sterilní. Z hlediska palynologie byly v
delších časových profilech zobrazených v sedimentech z vrtných jader patrné periodické
sekvence s bohatým obsahem palynomorf (místy s patrnými pseudomorfózami po
mikrokrystalech pyritu), střídající se s faciemi s naprostou převahou pylových zrn konifer
doprovázenými cystami mořských dinofyt s pouze výjimečně dochovanými dalšími
palynomorfami. Tato sekvence bývá následována sedimenty palynologicky sterilními,
případně sedimentací vápencových těles. K akumulaci pylových zrn konifer může docházet
v důsledku ekologických i tafonomických jevů. Většina pylových zrn konifer má pylová
zrna přizpůsobená přenosu větrem (vzdušné vaky). Hromadí se proto v mořských
sedimetech vzálených od pobřeží, kam se pylová zrna hmyzosnubných rostlin netransportují.
Mohou být navíc doneseny z morfologicky vyvýšených terénů, kde mají některé rody areál
původního výskytu (Tsuga, Cedrus, Picea, Ketelleria). Pylová zrna konifer jsou rovněž
odolnější vůči oxidickým podmínkám v mořské vodě i v sedimentu (Kwiatkowski, Lubliner
Mianowska 1957, Havinga 1964, 1967, Brooks 1971, Hopkins, Mc Carthy 2002). Hojnost a
diversita palynomorf v sedimentech je v přímé závislosti na oxidačně - redukčním potenciálu
(Martin, Drew 1970, Heusser 1978). Akumulace konifer je typická pro oligotrofní podmínky
mořské pánve a zvyšující se klimatické výkyvy. Tato klimatická nestabilita (cyklické změny
teplot, a humidity) je předpokládaným počátkem středně miocenní klimatické přechodné
fáze (MMCT) (Doláková et al. 2014, Holcová et al. 2015, Nehyba et al. 2016).
Výsledkem komplexního výzkumu badenských sedimetů je představa režimu pánve
ve vzájemných vztazích s podmínkami na přilehlé pevnině (Holcová et al. 2015, Nehyba et
al. 2016). Změna režimu v okrajových částech pánve byla v zóně MNN-5 zaznamenána v
časovém intervalu mezi posledním výskytem (LO) druhu Helicosphaera waltrans (14,36
mil. let) a posledním výskytem druhu Sphenolithus heteromorphus (Ca- nanoplankton).
V intervalu pod LO H. waltrans vznikaly ve studovaném území pouze siliciklastické
sedimenty, režim mořské vody byl více eutrofní - se značným obsahem živin, klima bylo
subtropické, poměrně vyrovnané, cyklicita pozorovatelná na mořských organizmech,
40
sedimentech i vegetaci byla nevýrazná. Palynospektra byla diversifikovaná a dobře
dochovaná (příklady vrtů Vyškov, Ivaň).
V intervalu nad LO H. waltrans byly patrné cyklické změny mořského režimu i
klimatu na kontinentu. Sedimentace siliciklastik byla doprovázena v okrajových částech
pánve vznikem karbonátových komplexů. Zaznamenány byly cyklické změny mořské
hladiny (transgrese, regrese) související s mírou cirkulace vody (dobré prokysličení/
stratifikace vodního sloupce), přínosu živin i teploty (zvyšování sezonality, zvyšování
salinity v připovrchových vodách související s aridizací klimatu). Každý cyklus začíná
poměrně náhlým oteplováním následovaným postupným ochlazováním. Kromě cyklických
změn v zastoupení teplomilných a akrtotercierních prvků spolu s místně zvýšeným
zastoupením heliofytních, suchomilných prvků (Olea, Buxus, Ephedra, Poaceae,
Asteraceae, Caryophyllaceae, Chenopodiaceae) se projevila cyklicita i ve stupni zachování
palynomorf. V periodách zvýšené cirkulace vody a dobrého prokysličení se vyskytovala
spektra s nadhodnocením Pinaceae. V tomto období byl rovněž na palynospektrech patrný
rozdíl v zastoupení vysokohorských prvků na opačných stranách mořské pánve. Na
lokalitách bližších k pasivnímu okraji českého masívu bylo zastoupení konifer poměrně
monotónní, bez výrazného poměru vysokohorských prvků. Tyto byly naopak častěji
přítomné na okraji pánve směrem k orogeneticky aktivnímu horstvu Západních Karpat, což
je dokumentováno na řadě lokalit např. Židlochovice, Lomnice, Rebešovice, Chrlice,
Otmarov, Opatovice, Oslavany (Doláková et al. 2014, Holcová et al. 2015, Nehyba et al.
2016).
Vznik karbonátového komplexu byl označen jako karbonátový event. (Holcová et al.
2015). Byl doprovázený snížením přínosu terestrického materiálu během transgrese, spojený
s oligotrofním režimem, rozšířením tzv. podmořských luk a aridizací klimatu.
Siliciklastické vrstvy materiálu uvnitř karbonátů byly palynologicky sterilní.
Podle těchto posledních výzkumů z karpatské předhlubně (Doláková et al. 2014,
Holcová et al. 2015, Nehyba et al. 2016) lze tedy biostratigraficky korelovat zónu vápnitého
nanoplanktonu MNN5 Zone nad LO (=Last Occurrence) Helicosphaera waltrans 4,6-13,9
mil. let s počátteční fází MMCT. Z dosavadních studií lze předpokládat, že cyklicita mořské
sedimentace byla vyvolána klimatickými změnami počátku MMCT.
41
4. Palynologie vybraných lokalit kvartéru
Vegetace kvartéru (obr. 8) je ovlivněna výraznými klimatickými výkyvy střídajících se
chladných a suchých období (glaciálů) a teplejších humidních fází (interglaciálů).
Vegetační pokryv krajiny, který se zobrazuje v pylových spektrech uchovaných
v sedimentech z archeologických lokalit, se vyznačuje různě intenzívním dopadem lidské
činnosti. V období pleistocénu a raného holocénu ještě nedocházelo k ovlivnění krajiny
lidskou činností v masivním měřítku. Základní charakter vegetace souvisel se změnami
přírodních podmínek. Pouze v těsném okolí lidských sídlišť byl rostlinný pokryv pozměněn
např. v důsledku intenzivněji ošlapávaných ploch nebo přibývající dotací dusíkatých látek
v důsledku životní činnosti. V mladších obdobích – počínaje kulturním obdobím neolitu –
začal člověk měnit krajinu zásadním způsobem zejména v důsledku zemědělství
(odlesňování a s ním spojený proces zvýšené půdní eroze a zvýšení intenzity povodňových
sedimentů, domestikace rostlin, extenzívní pěstování kulturních plodin, šíření plevelů,
změny původních areálů). Základní problematiku změn kvartérní vegetace v čase shrnují
v evropském měřítku zejména práce Firbas 1949, Lang 1994, Litt et al. 2008, Ložek 2007.
Problematikou vývoje vegetace z hlediska palynologie na území ČR se zabývali např.
Rybníčková (1974, 1985), Jankovská (1971, 1987), Svobodová (1990, 1991). V současné
době je studium změn vegetace (zejména od období konce posledního glaciálu) velmi
aktuální v souvislosti s porozuměním současnému vývoji a věnuje se mu celá řada mladých
specialistů.
V palynologických společenstvech z kvartérních sedimentů se pro zařazení
jednotlivých taxonů používá botanický systém.
42
Obr. 8. Stratigrafie kvartéru (Musil in Přichystal et al. 1996)
4a) Jeskynní sedimenty
Palynologická studia jeskynních sedimentů jsou ztížena tafonomickými problémy
v uchování a transportu palynomorf. Proto bylo těmto typům sedimentů na území ČR
věnováno pouze málo prací. Palynologickým studiem jeskynních sedimentů se do konce
devadesátých let zabývala pouze Svobodová (Svobodová 1988, 1992, Svobodová in Seitl et
al. 1986, Svobodová in Svoboda 1991). Jednalo se však vždy o sedimenty jeskynních vchodů
a otevřených prostor komunikujících s povrchem.
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Hlavním problémem studia palynologie ve vnitřních částech jeskyní je, že rostlinné
zbytky zde nejsou uloženy v blízkosti původního stanoviště, ale jsou do jeskynních prostor
druhotně transportovány (prosakující voda, spolu se sedimenty nebo v srsti a potravě
živočichů). Jedná se proto vždy o oryktocenozu vzniklou smíšením zbytků rostlin z různých
míst z povrchu jeskynních systémů. Často dochází i k redepozici a smíšení palynomorf ze
sedimentů uložených v různých časových obdobích. V sedimentech moravských krasových
oblastí dochází nejčastěji ke smíšení kvartérních a terciérních palynomorf. Značná část
rostlinných taxonů se ovšem na našem území vyskytovala v obou těchto útvarech (např.
borovice, jilm, olše, dub, lípa ale i byliny jako trávy, složnokvěté nebo i některé druhy
kapradin). Jejich přítomnost a rovněž množství v palynospektrech jsou přitom velmi
důležité pro stanovení klimatostratigrafie. Během transportu navíc dochází ke korozi a
degradaci zrn, a tudíž ani stav zachování a fosilizace nemusí být spolehlivým rozlišovacím
znakem. Tento fakt velmi ztěžuje přesné a jednoznačné stanovení stáří sedimentů, ale i
interpretace vegetace na povrchu jeskynních systémů. Jednou z možností odlišení různě
starých palynomorf představuje studium ve fluorescenčním mikroskopu (viz kap. 2b)
Dalším problémem typickým pro jeskynní palynofacie je selekce palynomorf
způsobená odlišnou rezistencí sporopylových obalů vůči transportu (mechanické interakce
s horninovými klasty) a chemismu prostředí během sedimentačních pochodů. Tento fakt se
projevuje zejména druhotným selektivním nahromaděním odolnějších pylových zrn a spor.
Je proto velmi obtížné stanovit, zda palynofacie s převahou určitých prvků ve spektrech
odpovídají původnímu ekologickému charakteru prostředí nebo vznikly v důsledku
mechanických a chemických procesů během sedimentace. Z moravských krasových území
byly zjištěny nadhodnocené akumulace různých pylových zrn a spor. Je evidentní, že
existuje několik faktorů ovlivňující tyto akumulace. Prvním z nich je klimaticky podmíněný
typ sedimentačního procesu. V příznivějších klimatických fázích se během transportu
akumulují zejména pylová zrna lípy a hladké monoletní spory kapradin čeledi Polypodiaceae
(dokumentováno např. z jeskyně Ochozské). Drobná pylová zrna podčeledi Asteroideae se
naopak kumulují v sedimentech vznikajících během chladného klimatu, např. lokality
Sloupsko-Šošůvské, Podhradní, Balcarka, Javoříčko (Doláková 2007, 2014)
Další zajímavý tafonomický jev popsali Navarro et al. (2001) z jeskynních sedimentů
Španělska. Autoři zjistili, že pylová zrna anemofilních rostlin (zejm. rod Pinus), která bývají
díky dobrému transportu větrem a velké pylové produkci v povrchových pylových spektrech
silně nadhodnocena, ubývají v palynospektrech z jeskynních sedimentů v závislosti na
vzdálenosti od jeskynního vchodu. Naopak pylová zrna rostlin zoofilních se směrem do
44
hloubky jeskyní na složení palynospekter podílejí mnohem častěji. Podle španělských
autorů se může jednat o transport způsobený živočichy (pylová zrna zoofilních rostlin bývají
morfologicky přizpůsobena ke snadnému zachycování na tělech živočichů).
V moravských jeskyních se tento jev podařilo dokumentovat v jeskyni Za hájovnou
v Javoříčském krasu (Doláková 2007), kde byla studována palynospektra ze dvou míst jedné
vrstvy. Sedimenty z profilu v Narozeninové chodbě, které byly situovány ve větší blízkosti
k původnímu jeskynnímu vchodu, obsahovaly převahu pylových zrn borovice a poměrně
hojné zastoupení lísky (větrosnubné). Naproti tomu oryktocenózy z téže vrstvy profilu
Kostnice II, který byl v pozici vzdálenejší, jsou o tato pylová zrna výrazně ochuzená a
naopak silně nadhodnocená o zástupce podčeledi Asteroideae. Spektrum bylo celkově
výrazně monotónnější.
Faktorem podílejícím se na zachování pylových zrn nepříznivě je vlhkost sedimentu,
protože v tomto prostředí se vyskytuje množství hub a bakterií, které rozkládají organickou
hmotu, a tudíž i pylová zrna (Navarro et al. 2001).
Z uvedených poznatků vyplývá, že pylová spektra z jeskynních sedimentů nelze jednoduše
porovnávat se standardními spektry platnými pro jednotlivá období kvartéru z povrchových
sedimentů (např. z rašelinišť). Jejich přesná determinace vyžaduje další studia, zejména
tafonomická, a neobejde se bez sedimentologického a paleontologického nebo
archeologického výzkumu. Podle studií autorů Carrión et al. (1999) lze za dostatečně
vypovídající o charakteru prostředí vně jeskynních sytémů považovat taková, kde bylo
determinováno více než 15 taxonů při minimálním počtu 200 pylových zrn a spor ve vzorku.
V rámci mé práce byla palynologická studia prováděna v sedimentech Moravského krasu
(jeskyně Ochozská, Balcarka, Sloupsko- Šošošůvská a Kůlna, Pod hradem, Dagmar, Barová,
Tereza, kaverny lomu Mokrá), Javoříčského krasu (Za Hájovnou) a Hranického krasu
(Hranická propast). Studované sedimenty byly kvartérního (holocén, pleistocén) i
terciérního stáří (Doláková 2007, Doláková 2014).
V Ochozské jeskyni byly studovány 2 profily, které měly obdobný charakter palynospekter.
Z palynologického hlediska se daly rozdělit do dvou částí. Ve spodní části profilů převládala
pylová zrna a spory vegetace chladné stepi (rody Helianthemum, Thalictrum, Selaginella
selagineloides, Saxifraga, Ephedra) spolu s teplotně nenáročnými dřevinami, odolnými vůči
chladu (Pinus cembra, Betula, Salix) a mokřinnými rostlinami (Cyperaceae, Potamogeton,
Botryococcus, Pediastrum). Nálezy cenobií zelené řasy druhu Pediastrum kawraiskyi
dokumentují existenci velmi chladného klimatu (Jankovská, Komárek 1982). Tyto části
profilů vznikaly s největší pravděpodobností v některé z chladných fází pozdního glaciálu.
45
Svrchní partie obou profilů byly typické jednotvárnými palynospektry s vysokou akumulací
pylových zrn lípy (Tilia) a hladkými monoletními sporami čeledi osladičovitých
Polypodiaceae. Přes tento vyselektovaný charakter palynospekter prokazují přítomná zrna
lípy charakter některého z teplejších klimatických období. Radiometrické datování sintrové
polohy v nadloží profilů detekovalo stáří vrstev sedimentů jako vyšší než 28 000 let (Kadlec
et al. 2000). Vylučuje tedy holocenní stáří těchto sedimentů s teplomilnějším charakterem
vegetace. Velmi obdobná palynospektra byla zjištěna ve stalagmitech z belgické jeskyně
Han - sur - Lesse, která byla datována do období 37 000 BP Bastin et al. (1987).
Profily, ve kterých byly patrné střídající se klimatické fáze posledního glaciálu (wisla)
byly zjištěny i v jeskyni Balcarka. Palynologická studia byla prováděna v zámci
záchranného archeologického výzkumu. Vchodové partie jeskyně jsou známy jako sídliště
magdalenských lovců (13 000 – 11000 BP) (Nerudová et al. 2010). Spodní části profilů
z vnitřních částí jeskynního systému obsahovaly oryktocenózy s převažujícími heliofytními
stepními rostlinami (Asteroideae, Artemisia, Poaceae, Ranunculaceae, Delphinium,
Chrysosplenium) a málo četnými dřevinami (Pinus, Betula, Alnus), které patrně odpovídají
chladné fázi pozdního glaciálu.
Nadložní sedimenty obsahovaly vyšší zastoupení dřevin s převahou lísky (Corylus),
hojnou borovicí (Pinus), břízou (Betula), olší (Alnus), lípou (Tilia) a sporadicky smrkem
(Picea). Tyto sedimenty pravděpodobně vznikaly během teplejších výkyvů posledního
glaciálu nebo během raného holocénu (boreál). K bližší charakterizaci by ovšem bylo třeba
dalších tafonomických, sedimentologických a archeologických výzkumů.
Palynologické studium spodních částí opěrného profilu ve vchodové části jeskyně
Kůlna poskytlo možnost srovnání s palynospektry vzorků odebíraných ve vnitřních částech
Sloupsko-Šošůvských jeskyní, které jsou s Kůlnou propojené. Opěrný profil v jeskyni Kůlna
byl podrobně zpracován z hlediska archeologického i paleontologického. Obsahuje
sedimenty, které se uložily v době od závěrečných fází saalského glaciálu (vrstva 14), přes
interglaciál eem (vrstvy 13,12,11a,c), poslední glaciál würm (wisla) (9b, 8a, 7a, b, c, 6) a
holocén (1-5) (Valoch et al. 1988). Jeho nejsvrchnější vrstvy palynologicky zpracovala
Svobodová (1988, 1992). Vrstvy 1- 4 klasifikovala jako holocenní, paleovegetace vrstvy 6
odpovídala chladné oscilaci pozdního glaciálu. Předmětem mého studia byly vrstvy 14 - 7.
Z palynologického hlediska bylo možné pozorovat klimatické oscilace, které se odrážely ve
vývoji vegetace. Palynospektra spodní části profilu dokumentovala zalesněnější krajinu
s mírnějším klimatem. Vrstva 14 indikovala oteplování v období závěru saalského (riss)
glaciálu. Vrstva 13 optimum a vrstva 11 zhoršení podmínek na konci interglaciálu eem.
46
Svrchní části profilu (vrstvy 9b - 7a, vrstva 7c byla radiometricky datována do období 45 000
BP) prokazovaly parkovitý charakter krajiny, kde se chladná stepní vegetace (Selaginella
selaginoides, Thalictrum, Ephedra, Botrychium) střídala s nehojnými lesními porosty s
nenáročnými dřevinami a častými mokřinami (Cyperaceae, Sphagnum). Mírně teplejší
charakter klimatu byl pozorován ve vrstvách 8a a 7c (sporadické nálezy taxonů Tilia, Acer,
Teucrium, Centaurea scabiosa). Mírné odchylky v charakteru vegetace a nálezech fauny
drobných savců mohly být způsobeny lokálními podmínkami v morfologicky členitém
povrchu Moravského Krasu. Vzorky z vnitřních partií Sloupsko – Šošůvských jeskyní
obsahovaly převahu bylinné stepní vegetace a byly korelovatelné s chladnějšími obdobími
z vrchní části profilu v Kůlně (würm až počátek holocénu). Tyto interpretace potvrdily i
nálezy části koster jeskynních medvědů (Ursus spelaeus).
Nejstarší studovaná palynospektra kvartérních sedimentů byla zjištěna v jeskyni Za
Hájovnou v Javoříčském krasu (Doláková 2007, 2014). Převážná většina studovaných
sedimentů byla paleontologicky i radiometricky datována jako interglaciál holstein (Musil
2005, Musil et al. 2014). Palynospektra byla studována z několika profilů převážně suťových
kuželů v různé vzdálenosti od původního jeskynního vchodu i z několika jeskynních
komínů. Palynologická studia potvrdila mírný charakter klimatu, optimální pro růst
smíšených lesů (Carpinus, Hedera, Acer, Tilia, Corylus). Ojediněle byla nalézána pylová
zrna rodů Pterocarya a Ilex, která představují prvky vegetace typické pro naše území
v neogénu. Tyto rody přežívají ve střední Evropě nejstarší kvartérní zalednění a opětně
migrují na naše území v teplejších obdobích. Představují vegetaci typickou pro klimatické
optimum holsteinského interglaciálu (Dyjakowska 1952, Vodičková-Kneblová 1961,
Břízová 1994, Bińka et al. 1997, Reille et al. 2000, Urban, Sierralta 2012). Jejich poslední
výskyt je dokumentován právě v interglaciálu holstein (Lang 1994, Litt et al. 2008, Roucoux
et al. 2008).
Jediná pylová spektra, která měla odlišný charakter, pocházela z jeskynního komínu.
Spektrum obsahovalo akumulaci pylových zrn podčeledi Asteroideae a nálezy vlhkomilných
prvků. V souhlasu s výsledky studia drobných savců vznikaly tyto sedimenty v
podmínkách chladnějšího klimatu a vegetační pokryv tvořila chladná step.
Změny palynospekter napomohly i interpretacím sedimentologickým, např. rozdělení
suťového kužele do dvou úrovní, které bylo podmíněno klimaticky. V profilech byla
pozorována i selekce, degradace a sekundární akumulace odolnějších pylových zrn
v důsledku chemickýcch, mechanických i mikrobiologických vlivů (viz výše).
47
4b) Archeologické lokality – vliv člověka na prostředí
Záchranný archeologický výzkum lokality Brno-Štýřice III zachytil v rámci střední
Evropy poměrně ojedinělý záznam lidského osídlení (epigravettien) v období krátce po
skončení posledního glaciálního maxima (LGM; 21 000 ± 2000 cal. BP). Interpretace
klimatu se opíraly o studia malakologie, palynologie a antrakologie (Nerudová et al. 2016).
Výsledky paleobotanické i malakologické analýzy dokumentovaly glaciální charakter
okolní krajiny s převládající světlomilnou vegetací charakteru travnaté stepi až keřovité
tundry (Poaceae, Helianthemum, Ephedra). Zjištěny byly otevřené lesní porosty břízy, vrby
a střemchy (Prunus padus), které byly podmíněny existencí příznivějšího mikroklimatu a
vlhčích stanovišť v okolí velkého vodního toku (Svratka). Lokalita představuje příklad
mikroklimaticky příznivějšího refugia pro lidské sídliště z období konce svrchního paleolitu.
Toto období je charakteristické absencí archeologické evidence ve značné části Evropy
(Feurdean et al. 2014, Heiri et al. (2014) potvrzující dekolonizaci obrovských, dříve
osídlených území vlivem extrémních klimatických podmínek předcházejících osídlení
magdalénienu (Nerudová et al. 2016).
V rámci vědeckovýzkumného záměru MSM0021622427 „Interdisciplinární centrum
výzkumů sociálních struktur pravěku až vrcholného středověku“ byly od roku 2005
z komplexního přírodovědného hlediska studovány zejména 2 významné archeologické
lokality: Těšetice a Pohansko.
Lokalita Těšetice je významné archeologické naleziště s polykulturním osídlením
zahrnujícím především sled hlavních neolitických kultur s dominantní starší fází moravské
malované keramiky (MMK), dále osadu únětického lidu a posléze i intenzívní osídlení
horákovské kultury doby halštatské (Podborský 1988). Od roku 1967 zde probíhá
systematický archeologický výzkum. Rozkládá se severozápadně od obce Těšetice a
jihozápadně od osady Kyjovice v nadmořské výšce 265-290 m v údolí potoka Únanovka.
Zatímco osada starší fáze MMK je situována na vyšších místech svahu, sídliště únětické a
halštatské je posunuto níže po svahu východním směrem. Na vrcholku táhlého návrší
lokality byl objeven rondel lidu s MMK (Podborský 1988).
Dřívější paleobotanický výzkum lokality prováděl Opravil (1961). Studován byl
především materiál uhlíků. Největší část vzorků pocházela z halštatských objektů, jen málo
z objektů doby bronzové a neolitu. Podle Opravila (1961) se vegetace těšetického halštatu
vyznačovala početným zastoupením světlomilných dřevin, hlavně křovin, rostoucích
48
obyčejně na volných prostranstvích, okrajích lesů anebo tvořících podrost ve světlých lesích.
Nejrozšířenější lesní společenstva byla svazu Quercion pubescentis. Vedle uhlíků byly
nalezeny v mazanici sídelních objektů otisky obilek a plev ovsa (Avena) (Opravil 1961).
V rámci výzkumného záměru bylo na lokalitě provedeno 5 mělkých vrtů (do10 m),
které většinou obsahovaly palynomorfy jen velmi sporadicky. Jediný vrt, který poskytl
bohaté palynologické asociace, byl vrt T4 situovaný v nivě říčky Únanovky cca 500 m jv.
od vlastní lokality. Výsledky tohoto vrtu byly propojeny s výsledky dalšího vrtu z nivy
Únanovky pod sídlištěm Staré zámky a společně publikovány (Petřík et al. 2014).
Sedimenty z obou vrtů v nivních sedimentech poskytly data z významné části období
holocénu (v rozmezí 5211-5008 cal. BC po současnost). Vrty by studovány za pomoci metod
sedimentologie, palynologie, makrozbytkové analýzy, malakozoologie, datování 14
C. Etapy
lidského osídlení byly konfrontovány s vývojem nivní sedimentace. Báze tvorby
sedimentace nivy byla zaznamenána na počátku neolitu. Okolní prostředí odpovídalo
smíšenému lesu s převahou dubu – Quercetum mixtum (Quercus, Tilia, Carpinus, Pinus),
s keřovitým podrostem (Corylus, Cornus). Bylinná složka byla tvořena zejména trávami a
ostřicemi (Poaceae, Cyperaceae). Na konci neolitu a počátku eneolitu se začínají v pylových
spektrech objevovat zrna obilovin. Klimatické změny během přechodu neolit/ eneolit a
vzrůstající intenzita lidské činnosti v souvislosti se zemědělstvím (odlesňování, orba,
zvyšující se pěstování kulturních plodin, zejména obilí) vedly i k výrazné akumulaci
sedimentů v nivě. Zvyšující se intenzita civilizačního impaktu byla patrná v průběhu celého
eneolitu. Ve spektrech se začínají objevovat rostliny typické pro ruderalizovaná stanoviště
(Artemisia, Chenopodium).
V následujícím období jsou změny v sedimentárním záznamu a vegetaci jen velmi
málo zřetelné. Od doby bronzové byla niva pokrytá zejména nitrofilní vegetací. Ačkoli na
studovaném území nejsou známé archeologické doklady pro období římské a stěhování
národů, v pylovém záznamu dokumentovaném C14 datováním (1320 BC - 1050 BC), jsou
z tohoto období zaznamenána zrna obilovin, která lidskou činnost v této době dokládají. Ve
vzorcích byla rovněž viditelná převaha bylin nad dřevinami, která by v tomto areálu bez
lidského zásahu nebyla pravděpodobná. Zvýšený podíl zaznamenala rovněž líska (Corylus),
vyskytující se v prosvětlených oblastech lesních okrajů, která mohla být i lidmi záměrně
šířená. Výrazná změna sedimentace i palynospekter byla viditelná od středověku. Podle
radiokarbonového datování byly tyto změny patrné od doby hradištní (230 - 410 AD).
Dřeviny ze spekter téměř vymizely, zastoupeny byly pouze v podobě lužních porostů
v zaplavovaném prostředí nivy (Alnus, Salix). Výrazné odlesnění se proto projevilo zejména
49
v sušších, pro pěstování plodin vhodných místech. Dominantní vegetace měla charakter
nitrifikovaných stanovišť (Sambucus, Chenopodiaceae, Cichorioideae, Urtica, Polygonum
aviculare). V sedimentačním záznamu od období vrcholného středověku do současnosti
byla intenzívní lidská činnost patrná z akcelerace eroze a akumulace v nivě doprovázená
zvýšenou koncentrací fosfátů (Petřík et al. 2014).
Lokalita Pohansko u Břeclavi představuje jedno z center raně středověkého
slovanského osídlení – Velkomoravské říše (9. století). Velkomoravské hradisko BřeclavPohansko
patří i v evropském měřítku k nejlépe prozkoumaným a zpracovaným raně
středověkým lokalitám svého druhu (Dresler, Macháček 2013). Systematické archeologické
výzkumy katedry prehistorie FF MU v Brně pod vedením prof. dr. Františka Kalouska a
prof. dr. Bořivoje Dostála zde byly zahájeny v roce 1958 (Vignatiová 1992). Pro vědecké
účely zde byla do roku 1990 odkryta, zdokumentována a vědecky vyhodnocena plocha větší
než 137 380 m2
(Macháček 2000).
Lokalita je situována v nadloží neogenních sedimentů vídeňské pánve. Kvartérní
sedimenty, ve kterých se nacházejí archeologické doklady, patří soutokové oblasti, v dnešní
době až několik kilometrů široké společné údolní nivě Dyje, Moravy a Kyjovky. Jedná se o
litologicky pestré říční (povodňové hlíny, jíly, fluviální písky, štěrky) a eolické sedimenty
(váté písky, spraše) a subfosilní půdy (Havlíček 2001, Macháček et al. 2007). Oblast je
jedním z nejníže situovaných areálů ČR. Nachází v nadmořské výšce 155-157 m. Kromě
velkomoravského osídlení je lokalita je známá jako stará sídelní oblast s archeologickými
doklady od mezolitu, přes neolit, eneolit, dobu bronzovou, dobu železnou reprezentovanou
hlavně obdobím laténu, dobu římskou a dobu stěhování národů, která zahrnuje nejmladší
předslovanské osídlení. V období Velkomoravské říše je známá jako jedno ze tří hlavních
sídelních center (Macháček 2005).
Širší okolí lokality byla centrem zájmu řady paleobotanických výzkumů zahrnujících
rekonstrukce vývoje vegetace údolní nivy Dyje a Moravy (Břízová and Havlíček (2002),
Jankovská et al. (2003), Opravil (1962, 1978, 1983b), Rybníček and Rybníčková (2001).
Rybníček a Rybníčková (2001) předpokládali během raného atlantiku v území nivy
vývoj travnatých subxerofylních doubrav a smíšených dubo-lipových lesů s jedlí na středně
vlhkých stanovištích. Vegetace byla intenzivně měněna vlivem odlesňování, pěstování
kulturních plodin a pastvy už během doby bronzové a starší doby železné.
Podle Ložka (2007) a Opravila (1999) způsobilo zvýšení srážek ve spodním atlantiku
zvýšení eroze a první zarovnávání níže položených nik v území údolní nivy. Od této doby
50
krajina postupně nabývá moderního charakteru. Podle Opravila (1978, 1983b, 1999, 2000)
měla vegetace v okolí raně středověkých hradišť oproti dnešnímu stavu vyšší diverzitu a jiné
kvantitativní proporce v důsledku vyšší morfologické rozrůzněnosti stanovišť. Vegetační
pokryv údolní nivy byl tvořen převážně lužními lesy a vlhkými loukami.
Jediné dřívější palynologické zpracování pocházející přímo ze sedimentů areálu
Pohanska publikovala Svobodová (1990). Ve studovaném profilu zdokumentovala zvyšující
se lidský impakt na vegetaci: odlesňování, přítomnost synantropních prvků (jako např.
Artemisia, Plantago lanceolata, Convolvulus arvensis, Centaurea cyanus, Viciaceae) a
kulturních rostlin (zejména obilniny Cerealia). Ve srovnání s dalším velkomoravským
centrem Mikulčice, které mělo charakter městské aglomerace s velkým zemědělským
zázemím, bylo Pohansko obklopeno smíšenými dubovými lesy. Bohužel, studovaný profil
neobsahoval archeologické záznamy, které by zpřesňovaly polohy s vlastním slovanským
osídlením.
V současné době představuje lokalita Pohansko jednu z nejdetailněji palynologicky
studovaných lokalit. Palynologická studia probíhala v rámci společného výzkumného
záměru a grantů Ústavu geologických věd PřF MU a Ústavu archeologie pravěku a
středověku FF. Bylo prostudováno 13 vrtů do 20m, 12 kopaných nebo zarážených sond, 2
bagrované profily, 11 profilů objekty, 21 vzorků z hrobových jam a více než 15 jednotlivých
vzorků.
Největší pozornost byla v součinnosti s archeology věnována profilům v rámci
archeologického řezu valem (profil před opevněnín, pod ním a na vnější straně, vzorky
z výplně mezi kameny valu). Nejnovější výsledky jsou v současné době odeslány nebo
připravovány do tisku v rámci interdisciplinárních výzkumů. Dosud publikované
palynologické výzkumy byly shrnuty zejména do prací Macháček et al. 2007 a Doláková
et al. 2010. Protože ne všechny profily a vrty poskytly kontinuální palynologická data, byla
část výsledků stratigraficky kompilována podle litologických charakteristik a C14 datování.
Nejspodnější části některých mělkých průzkumných vrtů zasáhly do
svrchnomiocenních sedimentů (pannon) vídeňské pánve, která tvoří podloží kvartérních
sedimentů. Jejich palynologický obsah byl diskutován v kapitole 3ab. V jejich nadloží byly
místy zachovány sedimenty pleistocenní v podobě hrubozrnných štěrků, které neposkytly
palynologické doklady. Největší část studovaných sedimentů patřila holocénu.
Nejstarší sedimenty datované metodou C14 pocházely z období mezolitu (14C in 8240
cal BP), nejmladší ze 16. století. Změny v proporcionálním zastoupení jednotlivých
rostlinných typů, které byly pozorovatelné v pylových diagramech, byly způsobené zejména
51
změnami povahy substrátu v závislosti na změnách humidity a rovněž lidskou aktivitou
(odlesňování, pěstování plodin, pastva).
Na bázi studovaných holocenních sedimentů byly palynologicky zdokumentovány
tafocenózy původních lesních porostů s nízkým ovlivněním lidskou činností. V pylových
spektrech převažovaly habrové doubravy s lípou (Quercus tvořil dominantní složku), které
se v čase i prostoru střídaly s lužními lesy (s převahou Alnus, méně Ulmus, Fraxinus, Salix
and Populus). Křoviny byly zastoupené méně, pravidelněji byla přítomná líska (Corylus).
Dřeviny ve spektrech převažovaly nad bylinami, ale krajina nebyla zcela zalesněná.
Nejstarší výrazný úbytek dřevin současně s nálezy pylových zrn obilnin (Cerealia) i
polních plevelů jako Polygonum aviculare (rdesno ptačí), (Centaurea cyanus) chrpa modrák
a druhotných antropogenních ukazatelů jako: jitrocel kopinatý (Plantago lanceolata) ,
šťovík kyselý (Rumex acetosella), chmel/konopí (Humulus/Cannabis) a merlíkovité
(Chenopodiaceae) prokázaly zemědělskou činnost už v období neolitu (S1: 127cm: 7050 BC
- 6450 BC). Protože byly tyto pylové asociace nalezeny ve vrstvách z těsného podloží
spodních povodňových sedimentů, jeví se tento výrazný lidský zásah do krajiny jako
pravděpodobná příčina zvýšeného výskytu povodní a akumulace povodňových hlín.
Výrazné lidské ovlivnění krajiny bylo dokumentováno rovn ěž v palynospekterch ze
sedimentů datovaných do období hallstattu (O1: 820 BC - 520 BC). Částečná rejuvenace
lesních porostů v nadložních sedimentech poukazuje na snížení intenzity lidského působení.
Tato obnova lesa byla potom následována nejvýraznějším odlesněním pozorovaným
v období existence velkomoravské aglomerace a vzniku jejího opevnění studovaného
zejména ve vzorcích z kultruní vrstvy vytyčené archeologickými nálezy. V nejmladších
povodňových sedimentech nad velkomoravskou vrstvou byla pozorována částečná
rejuvenace dřevin v podobě zvýšení nálezů zejména pionýrských porostů borovic a břízy
(Pinus, Betula). Došlo tedy ke snížení lidského impaktu.
V rámci palynologického studia na Pohansku bylo zjištěno několik zajímavých
problematik. V pylových spektrech se poměrně často objevovala pylová zrna ořešáku
(Juglans). Tato dřevina není na našem území původní. Místní nabohacení v pylových
spektrech umožňuje předpoklad záměrného pěstování, i když prozatím nebylo doloženo
makrobotanickými nálezy. Existence palynomorf vodních a vlhkomilných rostlin vně a
v těsném podloží opevnění a naopak jejich absence ve stejně starých sedimentech z vnitřní
strany valu mohou poukazovat na možnos, že opevnění fungovalo jako ochrana nejen proti
úočníkům, ale jako ochrana při povodních. Podporu pro toto tvrzení přinesl samotný řez
valem, kdy během intenzivních jarních povodních v r. 2007 došlo k zatopení velké části
52
vnitřní strany plochy hradiska. Výskyt pylových zrn habru (Carpinus) v sedimentech
datovaných do raného Atlantiku indikuje jeho dřívější šíření na území jižní Moravy, než je
tradičně předpokládáno pro území ČR jako celku (Firbas 1949), a podporuje tak interpretace
Opravila (1983a) a Rybníčkové (1985). Redepozice těchto pylových zrn z miocenních
sedimentů byly vyloučeny na základě studia autofluorescence.
53
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Přílohy:
Seznam přiložených publikací:
1. Bruch A. A., Utescher T., Alcade Olivares C., Doláková N., Ivanov D., Mosbruger
V. (2004): Middle and Late Miocene spatial temperature patterns and gradients in
Europe – preliminary results based on paleobotanical climate reconstructions. –
Cour.forsch.-Inst. Senckenberg, 249: 15-27.
Použito pro kapitolu: 3a
2. Doláková N. (2004): Discussion on some Thermophile palynomorphs from the
Miocene sediments in the Carpathian Foredeep (Czech Republic) and Modry Kamen
basin (Slovakia). – Acta Paleobotanica 44 (1), 79-85.
Použito pro kapitoly: 2, 3a
3. Doláková, N. (2007): Palynological studies in the Cave sediments from the
Moravian, Javoříčko and Hranice Karsts - Czech Republic. - Scripta Fac. Sci. Natur.
Universitatis Masarykianae Brunensis. 2007, Geology, 35, 47-53
Použito pro kapitoly: 2 4a
4. Doláková, N. (2014): Palynological analysis of Sediments from Za Hájovnou Cave.
- Acta Musei Nationalis Pragae, Series B – Historia Naturalis, 70 (1–2), 35–42.
Použito pro kapitoly: 2, 4a
5. Doláková N., Burešová A. (2007): Use of fluorescent microscopy in the study of
redeposited palynomorphs in the cave and marine sediments of Moravia (Czech
Republic). - Acta Palaeobotanica, 47, 1, 275-279.
Použito pro kapitoly: 2,4ab
6. Doláková N., Hladilová Š., Nehyba S., (1999): Development of Sedimentation,
molluscs and palynospectra in the Lower Miocene of the south-western Part of the
Carpathian Foredeep in Moravia (Czech Republic). - Acta Palaeobotanica, Suppl.
No. 2, W. Szafer Institute of Botany, Polish Academy of Sciences, Krakow, 269-278.
Použito pro kapitoly: 2,3a
7. Doláková N., Holcová K., Nehyba S., Hladilová Š., Brzobohatý R., Zagoršek K.
Hrabovský J., Seko M., Utescher T. (2014): The Badenian parastratotype at
Židlochovice from the perspective of the multiproxy study. - Neues Jahrbuch für
Geologie und Paläontologie, Abhandlungen, 271, 2, 169-201.
Použito pro kapitoly: 2,3a
8. Doláková N., Kováčová M. (2008): Pannonian vegetation from the northern part of
Vienna basin. - Acta Musei Nationalis Pragae Series B - Historia Naturalis 64, 2–4,
163–171
Použito pro kapitolu: 3a
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9. Doláková, N., Roszková, A. Přichystal, A. (2010): Palynology and Natural
Environment in the Pannonian to Holocene Sediments of the Early Medieval Centre
Pohansko near Breclav (Czech Republic). - Journal of Archaeological Science,
USA/UK : Elsevier, 37, 10, 2538-2550.
Použito pro kapitoly: 2, 4b
10. Doláková N., Kováčová M., Basistová, P. (2011): Badenian (Langhian – Early
Serravallian) palynoflora from the Carpathian Foredeep and Vienna Basin (Czech
and Slovak Republics). - Acta Musei Nationalis Pragae, Series B - Historia Naturalis
67 ( 1–2): 51-59.
Použito pro kapitoly: 2,3a
11. Doláková N., Slamková M. (2003): Palynological characteristics of Karpatian
sediments. - In: Brzobohatý R., Cicha I., Kováč. M., Rögl F. (Eds.) : The Karpatian
a lower Miocene stage of the Central Paratethys. Masaryk University Brno. 325-346.
Použito pro kapitoly: 2,3a
12. Holcová K., Hrabovský J., Nehyba S., Hladilová Š., Doláková N., Demeny A
(2015): The Langhian (Middle Badenian) carbonate production event in the
Moravian part of the Carpathian Foredeep (Central Paratethys): a multiproxy record
. - Facies, 61, 1, Article Number: 419
Použito pro kapitolu: 3b
13. Kováčová M., Doláková N., Kováč M. (2011): Miocene vegetation pattern and
climate change in the Northwestern Central Paratethys domain (Czech and Slovak
Republic). - Geologica Carpathica, 62, 3, 251—266
Použito pro kapitolu: 3a
14. Kvaček Z., Kováč M., Kovar-Eder J., Doláková N., Jechorek H., Parashiv V.,
Kováčová M., Sliva, Ľ. (2006): Miocene evolution of landscape and vegetation in
the Central Paratethys. - Geologica Carpathica, 57, 4, 295-310.
Použito pro kapitolu: 3
15. Nehyba S., Holcová K., Gedl P., Doláková N. (2016): The Lower Badenian
transgressive-regressive cycles – a case study from Oslavany (Carpathian Foredeep,
Czech Republic). - Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen,
Stuttgart, 279, 2, 209-238.
Použito pro kapitolu: 3b
16. Nerudová Z., Doláková, N., Novák, J. (2016): New information augmenting the
picture of local environment of the LGM/LGT in the context of the Middle Danube
Region. - The Holocene, 26(9), 1345–1354.
Použito pro kapitolu: 4b
69
17. Petřík J., Petr L., Šabatová K., Doláková N., Lukšíková H., Dohnalová A.,
Koptíková L., Blaško D. (2015): Reflections of prehistoric and Medieval human
activities in floodplain deposits of the Únanovka stream, South Moravia, Czech
Republic. - Zeitschrift für Geomorphologie, 59, 3, 393-41.
Použito pro kapitolu: 4b
View publication statsView publication stats
Acta Palaeobotanica 44(1): 79–85, 2004
Discussion of some thermophile palynomorphs
from the Miocene sediments
in the Carpathian Foredeep (Czech Republic)
and Modry´ Kamen basin (Slovakia)*
NELA DOLÁKOVÁ
Institute of Geological Sciences, Faculty of Science, Masaryk University, Kotlárˇská 2, 611 37 Brno, Czech Republic;
e-mail: nela@sci.muni.cz
Received 14 March 2003; accepted for publication 16 February 2004
ABSTRACT. The sediments from the southern part of the Carpathian Foredeep in Moravia (Czech Republic) and
the Modry´ Kamen basin from Slovakia were studied. The sediments are of Eggenburgian to Lower Badenian
ages and of marine or brackish origin. The marine facies rapidly changes with lagoonal and deltaic facies during
the Eggenburgian–Ottnangian. The palynospectra have warm-subtropical character. Saltmarsh vegetation (37%
Chenopodiaceae) interchanged with swamp vegetation. In the Karpatian palynospectra marsh facies occur more
uniformly. Marked azonal associations, for example marsh palm forest, riparian forest or the associations with
Taxodiaceae and Tiliaceae (Craigia), are typical. The Ottnangian–Karpatian palynospectra from the Modry´
Kamen basin were typical in the occurrence of pollen of Pentapollenites sp. The studied Lower Badenian sediments
represent the fully marine conditions. The palynospectra were poor in pollen and spores (except Pinaceae)
and rich in Dinoflagellata. At this time the climate was not extremely warm.
KEY WORDS: palynology, Miocene, Carpathian Foredeep, Czech Republic
INTRODUCTION
The sediments from the southern part of
the Carpathian Foredeep in Moravia (Czech
Republic) were studied palynologically. Palynomorphs
have been described from many boreholes
and outcrops with the sediments of the
Eggenburgian to Badenian age. Palynological
studies were compared primarily with sedimentological
and palaeontological investigations.
The studies were directed firstly on the
relationship between the terrestrial and
marine ecosystems. Palynological results were
compared with the ones from the Modry´
Kamen basin in the southern part of the Slovak
Republic.
RESULTS
The sediments are of Eggenburgian to the
Lower Badenian age, and are marine or brackish
in origin. As a whole, the palynospectra
indicate a warm-subtropical character of
climate. Sapotaceae, Palmae, Engelhardia,
Platycarya, Castaneoideae, Tricolporopollenites
liblarensis (Th.) Th. & Pf., thermophile
oaks, Lygodium, and Pteridaceae are frequent.
Symplocos, Reevesia, Cornaceae, Parthenocissus,
Tricolporopollenites pseudocingulum
(Pot.) Th. & Pf., and families Araliaceae and
Rutaceae occur regularly but in lower quantities.
Alangium and Neogenisporis neogenicus
Krutzsch are represented by sporadic occurrences
only. Arctotertiary elements are slightly
less frequent. Practically all of the palynospectra
are strongly influenced by sedimentological
* This study was supported by the Grant Agency of the
Czech Republic, grant no. 05/00/0550.
This paper was presented on the EEDEN Workshop, Kraków
28 June – 1 July 2002
facies. This fact is reflected in the proportional
changes between the palaeotropic and arctotertiary
elements, and it is very difficult to
specify any trends in the climate development
from these data. Higher percentages of the
arctotertiary elements were observed locally,
for example:
– Chenopodiaceae, Oleaceae, Ericaceae,
Salix, Potamogeton, Ulmaceae, Taxodiaceae –
in the sediments of the Eggenburgian–Ottnan-
gian,
– Alnus, Ulmaceae, Osmunda, Polypodiaceae,
Lythraceae, Sparganium – in the Karpatian
sediments.
In the pollen diagrams of the studied
Miocene sediments, no explicit climatic
changes are visible. The author cannot specify
at the present time if the observed changes are
influenced climatologically or caused by the
development of the sedimentary basin and adjacent
areas.
The environment of the studied part of the
Carpathian Foredeep was extraordinarily
variable during the Eggenburgian–Ottnangian.
The marine transgression penetrated the
sea coast with highly differentiated relief configurations.
The marine facies interchanged
rapidly with those of lagoons and deltas. Sediments
and molluscs show rapid changes in salinity,
dynamics and depths, light and evaporation.
The palynospectra reflected many of
these changes (Nehyba et al. 1997).
Due to oscillations in salinity and occasional
higher levels of evaporation halophilous
vegetation grew on the coast (up to 37%
Chenopodiaceae, Pl. 1, fig. 9). In some places
these were accompanied with higher number
of Ephedra and Buxaceae. Ericaceae are frequent
in the facies manifesting a low salinity.
Pollen grains of the species cf. Monocirculipollis
zahnaensis Krutzsch (Pl.1, figs 1–4) in
one case even in pollen conglomerate, which
most probably contradicts the redeposition
from older sediments, were typical for the
sediments of the Eggenburgian age. These pollen
were found in all the Eggenburgian boreholes
and they were absent in younger sediments.
Krutzsch (1966) described these pollen
types from the Palaeocene and Eocene sediments
and he considered them as extinct
members of the family Buxaceae. Similar pollen
types were found also in the Miocene sediments
from Turkey (Nurdan Yavus, pers.
comm.). According to data in the literature, the
pollen type seem to be similar to some members
of the families Caryophyllaceae or Amaranthaceae.
It is necessary to test this opinion
with SEM observations on the detailed features
of the pollen.
Even the presence of small tri- or tetracolporate
grains of Rutaceae is typical for some
localities. A large amount of Platanus pollen
was also observed in some samples.
Saltmarsh vegetation developed in time and
space to the various growth stages of the
swamp and riparian vegetation. For example
facies with Myrica pollen (Pl. 1, fig. 13) overlapping
40%, or Taxodiaceae approaching
values of 50% are visible. The freshwater flora
with Nelumbo, Potamogeton, Sparganium, and
Cyperaceae has been ascertained. Lygodium
(Pl.1, fig.14) was also frequent, in one locality
its frequency reached more than 5%.
Later on, the character of the coast development
changed, with the relief becomming flatter.
The transgression stage follows in the Ottnangian,
the sea probably penetrated further
to the north into the area. The sediments
include the so called Rzehakia beds (Nehyba et
al. 1997). Facies with moorland and marginal
swamp appeared in the palynospectra. Taxodiaceae
are also frequent. The climate at this
time appears to be more humid. Pollen spectra
contain a larger amount of spores of thermophile
ferns as Lygodium, Pteridaceae, Gleicheniaceae
together with Riccia and Selaginella
(Pl. 1, fig. 10). Ilex is relatively plentiful. The
higher representation of Symplocos pollen is
interesting in the samples of Ottnangian age.
Striking members in some palynospectra
are the higher proportions of Selaginella (up to
6%), their occurrence being visible in the Eggenburgian/Ottnangian
and Ottnangian/Karpatian
border sediments. They are accompanied
by an increased frequency of other
pteridophytes. The humid climate was probably
connected with the transgression.
The Karpatian sedimentation began by
gradual transgression on the relatively flat
coast, which was connected with anoxic conditions.
The frequent alteration of palynomorphs
caused by precipitation and growth of pyrite is
visible. Later on, fully marine conditions with
schlieric sediments developed. In the Karpatian
palynospectra the marsh facies occur
more uniformly than in the Eggenburgian–
Ottnangian. Pollen and spores also confirm
a warm to warm-temperate climate.
80
The markedly azonal associations, for
example marsh palm forest with Palmae (Pl. 2,
figs 9,10), Poaceae, Lygodium, Sparganium,
and Potamogeton; riparian forest with Alnus,
(Pl. 2, fig. 4), Myricaceae (Pl. 2, fig. 3), Lythraceae
(Pl. 2, figs 1,2), or Selaginella (Pl. 2, fig. 13)
are frequent in Karpatian sediments. The associations
with Taxodiaceae, Intratriporopollenites
insculptus Mai (Pl.2, figs 6,7) and Pteridaceae
(up to 10%) are typical. According to
recent investigations (Kvacˇek et al. 2002) Intratriporopollenites
insculptus belongs to the
genus Craigia (Malvaceae–Tilioideae). Very
similar associations were also described from
the lower Miocene of Northern Bohemia (Konzalová
1976). These pollen types were found
also in the Eggenburgian sediments of the
Carpathian Foredeep. In all the cases they are
accompanied with high amount of other hygrophilous
elements (except the ones mentioned
above also Ericaceae and Salix).
Assotiations of markedly colder climatic
conditions, known from the adjacent areas
(e.g. Slovakia, Austria) from the end of the
Ottnangian and the lower Karpatian (Hochuli
1978, Planderová 1990) have not been found.
Proportional changes between palaeotropical
and arctotertiary elements seems to be
influenced by facies in all of the samples
studied.
The schlieric sediments, representing
marine conditions, contain a large amount of
Pinaceae and an increasing amount of Oleaceae
(Pl. 2, figs 8,12). Sapotaceae have not
been found in the uppermost studied Karpatian
sediments. Arctotertiary elements
increased slightly. However, there are not sufficient
data to enable a more exact interpretation
of this as representing a colder climate in-
terval.
Conglomerations of pollen (Monocirculipollis,
Chenopodiaceae, Platanus, Oenotheraceae,
Myricaceae, Alnus, and others) were found in
some facies from the whole of the Lower
Miocene. Their presence confirms the low-flow
water dynamics and short transport distances
to the place of sedimentation.
The studied sediments from the Modry´
Kamen basin include the Rzehakia beds and
the schlieric layers (Ottnangian–Karpatian).
The palynospectra are similar to the ones from
the Carpathian Foredeep. The biggest difference
is based on the typical occurence of
pollen of the formal genus Pentapollenites
(Pl. 2 fig. 14–16) which after Krutzsch (1962)
is characteristic for Palaeogene. These pollen
have also been described from Miocene sediments
by Planderová (1990) and Nagy (1985).
Planderová interprets these types of pollen as
typical for the coal facies of the Slovakian
Lower Miocene. However, the author has not
observed these pollen types in the Carpathian
Foredeep. According to the recent literature
(Reille 1995) part of our material is very close
to the genus Haplophyllum (Rutaceae). Hofmann
and Zetter (2001) described this pollen
from the Palaeocene/Eocene of Austria and
considered them belonging within the Simaru-
baceae.
The studied Lower Badenian sediments
represent the development of fully marine conditions
with a high oxygen content. The palynospectra
are generally poor in pollen and
spores (except for Pinaceae), and are comparatively
rich in Dinoflagellata. The tapeta of foraminifers
have also been identified. It is probable
that the conifer pollen accumulated in
these marine sediments due to their wind dispersal
properties.
The environment on the seashore was probably
rather wet as noted by the presence of
spores of Fungi, Polypodiaceae, pollen of Alnus
and Ulmus and ranging up to swampy as represented
by the pollen of Taxodiaceae and
Myricaceae. There are visible only slight
changes in the pollen diagrams between the
Karpatian and Lower Badenian pollenspectra.
In the Badenian a decreasing number of some
thermophilous members, such as Engelhardia,
Platycarya, Tricolpopollenites henrici (Pot.) Th.
& Pf. and T. microhenrici (Pot.) Th. & Pf., and
Castanoideae are observed. Oleaceae were also
less frequent, although Taxodiaceae regularly
have a higher representation while Alnus has
steadily lower percentages. In the Badenian
palynospectra the higher differentiation of the
pollen of thermophilous forms of oaks such as
Tricolpopollenites henrici (Pot.) Th. & Pf. and
T. microhenrici (Pot.) Th. & Pf. or forms probably
similar to the Quercus ilex as well as deciduous
ones, are observed (Pl. 2, figs 18–19).
The present author cannot currently specify
if the changes observed are influenced climatologically
or are caused by the development of
the sedimentation basin and its adjacent
areas.
81
REFERENCES
HOCHULI P.A. 1978. Palynologische Untersuchungen
im Oligozän und Untermiozän der Zentralen und
Westlichen Paratethys. Beitr. Paläont Öster., 4:
1–132.
HOFMANN Ch.Ch. & ZETTER R. 2001. Palynological
investigation of the Krappfeld area, Palaeocene/Eocene,
Carinthia (Austria). Palaeontographica,
B, 259 (1–6): 47–64.
KONZALOVÁ M. 1976. Microbotanical (palynological)
research of the lower Miocene of Northern Bohemia.
Rozpr. Cˇ es. Akad. Veˇd, 86 (12): 1–75.
KRUTZSCH W. 1962. Mikropaläontologische (sporenpaläontologische)
Untersuchungen in der Braunkohle
des Geiseltales. II. Die Formspezies der
Pollengattung Pentapollenites Krutzsch 1958. Paläont.
Abh., 1(2): 75–103.
KRUTZSCH W. 1966. Zur Kenntnis der präquartären
periporaten Pollenformen. Geologie, 15(55): 16–71.
KVACˇ EK Z., MANCHESTER S.R., ZETTER R. & PINGEN
M. 2002. Fruits and seeds of Craigia bronni
(Malvaceae-Tilioideae) and associated flower
buds from the late Miocene Inden Formation,
Lower Rhine Basin, Germany. Rev. Palaeob.
Palyn., 119: 311–324.
NAGY E. 1985. Sporomorphs of the Neogene in Hungary.
Geologica Hungarica, ser. Palaeont., 47:
1–470.
NEHYBA S., HLADILOVÁ Š. & DOLÁKOVÁ N. 1997.
Sedimentary evolution and changes of fossil assemblages
in the SW part of the Carpathian Foredeep
in Moravia during the Lower Miocene. In:
Hladilová Š. (ed.) Dynamika vztahu˚ marinního a
kontinentálního prostrˇedí. Sborník prˇíspeˇvku˚.
Grantovy´ projekt GACˇ R 205/95/1211. Masarykova
univerzita: 40–58. Brno. (In Czech with English
summary).
PLANDEROVÁ E. 1990. Miocene Microflora of Slovak
Central Paratethys and its Biostratigraphical
Significance. Geol. Inst. D. Štúra, Bratislava.
REILLE M. 1995. Pollen et Spores d’ Europe et d’ Afrique
du Nord, (I,II). Marseille.
Plate 1
Sporomorphs of Eggenburgian – Ottnangian
× 1000, except figs 3,4 and 13
1–4. Monocirculipollis zahnaensis Krutzsch, Cˇ ejkovice 176.8 m, Eggenburgian–Ottnangian
3. SEM × 2300
4. SEM × 5500
5. Reevesiapollis triangulus (Mamczar) Krutzsch, Líšenˇ, Ottnangian
6. Sapotaceoidaepollenites sp., Miroslav 78.4 m, Eggenburgian–Ottnangian
7, 8. Rutaceaepollenites sp., Šafov 12, 17.5 m
9. Chenopodipollis multiplex (Weyl. & Pf.) Krutzsch, pollen conglomerate, Miroslav 78.4 m,
Eggenburgian–Ottnangian
10. Echinatisporis miocenicus Krutzsch & Sontag in Krutzsch, Trboušany 65.8 m, Eggenburgian–Ottnangian
11. Symplocoipollenites vestibulum (Potonié) Potonié, Líšenˇ , Ottnangian
12. Potamogetonpollenites sp., Líšenˇ , Ottnangian
13. Myricipites sp., pollen conglomerate, Trboušany 49.7 m, Eggenburgian–Ottnangian, × 500
14. Leiotriletes maxoides maximus (Pflug) Krutzsch, Trboušany 49.7 m, Eggenburgian–Ottnangian
PLATES
82
Plate 1 83
N. Doláková
Acta Palaeobot. 44(1)
Plate 2
Sporomorphs of Ottnangian–Karpatian–Badenian
× 1000
1,2. Lythraceaepollenites sp., Ždánice 67, 780–785 m
3. Myricipites rurensis (Pf.& Th.) Nagy, Ždánice 67, 780–785 m, Karpatian
4. Alnipollenites verus Potonié, Ždánice 67, 785–790 m, Karpatian
5. Leiotriletes wolffii wolffii Krutzsch, Nosislav 3, 323 m, Karpatian
6,7. Intratriporopollenites insculptus Mai, Ždánice 68, 815–820 m, Karpatian
8,12. Oleoidearumpollenites microreticulatus (Th. & Pf.) Ziembin´ska-Tworzydło, Nosislav 3, 280.8 m, Karpatian
9,10. Sabalpollenites areolatus (Potonié) Potonié, Ždánice 67, 795–800 m, Karpatian
11. Potamogetonpollenites sp. Ždánice 67, 795–800 m, Karpatian
13. Echinatisporis miocenicus Krutzsch & Sontag in Krutzsch, Nosislav 3, 343 m, Karpatian
14. Pentapollenites fsp. 1, Modry´ Kamen Basin, N91, 345 m, Ottnangian–Karpatian
15, 16. Pentapollenites fsp. 2, Modry´ Kamen Basin, N91, 345 m, Ottnangian–Karpatian
17. Marine Dinophyta, Židlochovice, Badenian
18. Quercoidites granulatus (Nagy) Słodkowska, Židlochovice, Badenian
19. Quercoidites sp. 1 – Quercus ilex type, Židlochovice, Badenian
20. Cercidiphyllites minimireticulatus (Trevisan) Ziembin´ska-Tworzydło, Židlochovice, Badenian
21. Platanipollis ipelensis (Pacltová) Grabowska, Lysice, Badenian
22. Polypodiaceoisporites corrutoratus Nagy, Židlochovice, Badenian
23. Inaperturopollenites hiatus (Potonié.) Th & Pf. – Glyptostrobus type, Ždánice 68, 815–820 m, Karpatian
84
Plate 2 85
N. Doláková
Acta Palaeobot. 44(1)
Introduction
Palynological study of sediments from Za Hájovnou Cave
(Javoříčko karst) was carried out within the framework of
a complex multiproxy study of this area headed by
Prof. R. Musil.
Results of earlier palynological studies of Moravian (the
eastern part of the Czech Republic) karstic areas were
published by Seitl et al. (1986), Svobodová (1988), Svoboda
(1991), Svobodová (1992), Doláková and Nehyba (1999),
Doláková (2002, 2004, 2005, 2007).
Pollen spectra from cave sediments are typified by
the absence of in situ plant remains. Palynomorphs are
transported into the caves together with sedimentary particles
by percolating water or through the activity of animals.
Selection, degradation and secondary accumulation of
various palynomorphs is clearly due to their different
resistances to chemical and mechanical processes, and
microbial attack during transport and sedimentation (Elsik
1971, Havinga 1971, Jankovská 1971, Draxler, 1992, Carrión
et al. 1999, Doláková and Nehyba 1999, Navarro et al.
2001, Doláková 2002, 2007). The mixing of different age
components – especially Quaternary and redeposited Tertiary
elements – is also well known (eg. Doláková and Nehyba
1999, Doláková 2002, 2007). This phenomenon causes
complications in interpretation of the original surface
vegetation. Comparison with the results from other
paleontological and geological methods is necessary.
Material and methods
About 50 samples of cave sediments from Za Hájovnou
Cave were studied from a palynological point of view. The
palynological samples were treated with HCl (20%), HF,
KOH and HCl (10%) and heavy liquid ZnCl2 (density =
2g/cm3
) for standard maceration. The omission of acetolyse
enabled clearer identification of pollen contamination eg by
percolating water. The final residue from each sample was
mounted in preparation for biological microscopy, and
diluted with glycerol.
The pollen diagram was calculated from the total of
a minimum of 100 determined pollen grains and spores
(minimally from 15 taxons) using the POLPAL programme
(Walanus and Nalepka 1999). Several plant types were
combined according to their ecological grouping (Carpinus +
Tilia, Quercus + Acer, herbs undif., herbs aquatic, flood plain
forest – Alnus, Ulmus, Fraxinus, Salix, ferns + Sphagnum).
The pollen diagram was separated into two parts due to
the over-representation of Asteraceae: in the left section was
the pollen sum (100%), excluding Asteraceae. The right
section showed the proportion of Asteraceae and was
calculated from the sum of all the determined pollen grains.
This form of presentation offers clearer visualisation of the
basic character of the vegetation changes.
A composite pollen diagram (Text-fig. 1) was constructed
on the basis of average representation of elements from single
layers (arithmetic mean of samples collected from several
places within 1 layer). Other diagrams (Text-figs 2a, b)
indicate differences among the palynospectra collected from
different spots within a single layer.
Results
The samples from profiles: ZH P-2 (Velikonoční jeskyně
(= Easter corridor): Komín I (= Chimney I), ZH P-5
(Vykopaná chodba (= Excavated corridor): Kostnice II
35
SBORNÍK NÁRODNÍHO MUZEA V PRAZE
Řada B – Přírodní vědy • sv. 70 • 2014 • čís. 1–2 • s. 35–42
ACTA MUSEI NATIONALIS PRAGAE
Series B – Historia Naturalis • vol. 70 • 2014 • no. 1–2 • pp. 35–42
PALYNOLOGICAL ANALYSIS OF SEDIMENTS FROM ZA HÁJOVNOU CAVE
NELA DOLÁKOVÁ
Masaryk University, Faculty of Sciences, Institute of Geological Sciences, Kotlářská 2, 611 37, Brno, the Czech Republic;
e-mail: nela@sci.muni.cz
Doláková, N. (2014): Palynological analysis of sediments from Za Hájovnou Cave, Javoříčko Karst. – Acta Mus. Nat. Pragae, Ser. B, Hist.
Nat, 70(1-2): 35–42, Praha. ISSN 1804-6479.
Abstract. The sediments of profiles ZH P-2, 5, 7, 8b, 9, 10 and 11 from Za Hájovnou Cave (Javoříčko Karst) were studied from a palynological
point of view. Most of layers (except layer 1c/ZH P-10, layer 7e /ZH P-2) were paleontologically dated as Holsteinian Interglacial. The palynospectra
confirmed the mild character of the climate during sediment deposition (Carpinus, Hedera, Acer, Tilia, Corylus, single Pterocarya
and Ilex). The proportion of individual elements as well as relationship between trees and herbs varied. Changes in number of morphologically
different pollen grains (primarily Pinaceae, Asteroideae) in correlated layers (2a, 2b/ ZH P-5 and 2,2b/ZH P-8b, and layer 4 – debris cone)
from profiles ZH P-10, ZH P-11and ZH P-8b were recorded in the direction towards the cave interior. This phenomenon is most likely related
to the resistance of the pollen grains to chemical and mechanical conditions during transport.
■ Javoříčko karst, Za Hájovnou Cave, Quaternary, Palynology
Received January 20, 2014
Issued October, 2014
DOI 10.14446/AMNP.2014.35
36
Text-fig. 1. Composite pollen diagram.
Text-fig. 2. Pollen diagram. a – layer 2b from ZH P-5 correlating with 2b of ZH P-8 nd 9; b – layer 4 (debris cone) of profiles ZH P-10
and ZH P-11.
(Charnelhouse II), ZH P-7 (Chodba naděje (=Corridor of
Hope)), ZH P-8b and 9 (Spojovací chodba/Narozeninová
chodba (=Connection Passage/Birthday corridor)), ZH P-10
and 11 (Narozeninová chodba) were analysed from
palynological point of view. Profile descriptions are presented
in the contribution by Musil (2014). The first palynological
study from Za Hájovnou Cave was published by Doláková
(2005, 2007). Not all the palynospectra contained a sufficient
number of pollen grains and spores for analysis. Some of the
samples were almost sterile, another part contained only
a small number of palynomorphs. Samples with the most
abundant palynospectra came from profiles ZH P-8b and 9.
Only layer 4 of profiles ZH P-10 and ZH P-11 and layers 2a
and 2b from profile ZH P-5 contained enough pollen grains
and spores to construct a pollen diagram.
Most of the palynologically studied sediments (except ZH
P-1, layer1c and ZH P-2, layer 7e) were dated as Holsteinian
Interglacial (Musil 2005, Lundberg et al. 2014).
Sample ZH P-10, layer 1c was assigned as a melange of
Holocene and older sediments (Musil 2014). It contained
only a small amount of pollen from Pinus, Asteraceae,
Daucaceae and Ranunculaceae.
The sample from layer 7e of ZH P-2 was the only one
from the Komín I profile in which palynomorphs were found.
There was a large over-estimation of small Asteroideae.
Pollen from Chrysosplenium/R.trichophylus type typical for
damp places had also accumulated here along with some
other herbs (Poaceae, Galium, Ranunculaceae). No pollen
from trees or large sized pollen grains were observed. This
oryctocoenoses provides evidence of mechanical selection
during transport with sediment flow. According to a personal
study of cave sediments (Balcarka – Doláková 2004, Pod
Hradem – personal data of the author) such palynospectra
could support the paleozoological interpretation of
development in a colder steppe environment (Musil 2005,
Ivanov 2005).
The palynospectra from other studied samples of layer 4
(debris cone) from profiles: ZH P-10 and 11, ZH P-5: layers
2a, 2b and the whole ZH P-8a similarly confirmed the mild
character of the climate during sediment deposition, by virtue
of the abundant wood elements (Carpinus, Tilia, Juglans,
Quercus, sporadic Acer, single Hedera, Pterocarya, Ilex -
Tab.1).
Pterocarya and Ilex are the surviving members of
Tertiary floras. The presence of Pterocarya continued to the
Holsteinian Interglacial in Europe. It dissapeared from the
palynological record during the Saalian complex Stage and
it is not known from any younger warmer phases (Lang 1994,
Litt et al. 2008, Roucoux et al. 2008).
The above mentioned plants created the typical
vegetation of the climatic optimum of the Holsteinian
Interglacial (Dyjakowska 1952, Vodičková-Kneblová 1961,
Břízová 1994, Lang 1994, Bińka et al. 1997, Reille et al.
2000, Urban and Sierralta 2012).
The general character of most studied pollen spectra is
quite similar, however the proportions of individual elements
fluctuate. Composite pollen diagrams indicate changes
among pollen spectra in recognised layers (Text-fig. 1). The
relationship between trees and herbs varied, herbs mostly
prevail. Dominance of arboreal pollen was observed in
layers: 2b and 5: 1.2–1.5 m and 9.5–10 m from ZH P-8b and
9, and in part of the debris cone: layer 4 upper. Pinus
sylvestris type or Corylus formed the highest proportion of
them. Elements of flood plain forest such as Alnus, Ulmus,
Fraxinus, Salix were mostly rare. Betula, Picea and ferns and
Sphagnum are observed continuously but at a low percentage.
Nonarboreal pollen dominated in most samples. Asteraceae
prevailed in these cases. Poaceae were also common. The
occurrence of other herbs was also recorded: Artemisia,
Brassicaceae, Campanula, Galium, Lamiaceae, Liliaceae,
Ranunculaceae, Delphinium type, Silenaceae, Urtica and
single specimens of other herbs. Hygrophilous herbs such as
Chrysosplenium/Ranunculus trichophylus type and Valeriana
were common. Pollen from aquatic flora (Sparganium,
Potamogeton) were rare. Ferns mostly represented by the
smooth spores of Polypodiaceae, rarely Polypodium vulgare
and Pteridium. Algae such as Botryococcus, Mougeotia and
a single Pediastrum were found locally. Algae were absent
in samples from the debris cone (layer 4).
The greatest difference in pollen picture was observed in
layer 5, the deepest studied layer: 9.5-10 m in profile ZH P-9.
Trees prevailed over herbs (67:33%) with the most abundant
being Corylus together with Carpinus, Tilia and Alnus.
Juglans also occurred here. In this layer was the lowest
proportions of Pinus and Asteroideae from all the samples.
Only Poaceae represented the more abundant herbs. This
palynospectrum probably represented the vegetation of
deciduous woodland with only a small admixture of conifers.
In the overlying samples there was an increased proportion
of Pinus, Asteroideae and other herbs and a decrease in
Corylus, Alnus and Tilia until they became absent at the
6.5–7.5 m level. It is difficult to decide whether this
phenomenon is a result only of climate deterioration or also
due to taphonomic reasons.
The highest percentage of Pinus (over 60%) together with
a minority of other trees was recorded in the upper part of
the debris cone (layer 4 upper). The pollen picture in the
lower part of the debris cone differs with Pinus representing
only 10% of the pollen found, Corylus over 25%, Carpinus
and Tilia also occured, but more herbs were common
(Text-fig. 1). This difference could indicate climatic
variations during deposition of the debris cone.
According to paleontological results, layers 2a and 2b
from ZH P-5 correlate with 2a and 2b from ZH P-8b. The
pollen diagram from these layers indicates a decreasing
number of Pinaceae and dominance of tree pollen, and
increasing number of herb pollen (mainly Asteroideae) in the
direction towards the cave interior (Doláková 2005)
(Text-fig. 2a). These facts are most likely related to the
resistance of the pollen exines to chemical and mechanical
conditions during transport. A similar phenomenon was
observed in layer 4 (the debris cone) of the profiles ZH P-10
and ZH P-11 (Text-fig. 2b). Differences in the pollen picture
were probably caused by mixing of sedimentary material
transported through the former cave entrance and near the
chimney. A difference in pollen record between samples of
layer 4 (the debris cone) collected from ZH P-10 and ZH
P-11 compared to ZH P-8b is evidence enough to indicate
that sedimentation of this layer occured over a longer time
span during slight climatic oscillation. Therefore the layer
was divided into 2 sublayers: layer 4 upper and the
underlying: layer 4 lower (Text-fig. 1).
37
Discussion
Navarro et al. (2001) established that the amount of the
anemophilous pollen (eg. Pinus) decreased and that the
amount of zoophilous pollen (Asteraceae/Cichorioideae)
increased in the direction from the cave entrance into the
inner parts. They presumed that the anemophilous pollen,
often overestimated in surface samples due to massive pollen
production and extensive flight range, decreased inside the
caves due to their mechanical properties. Conversely, the
anemophilous pollen grains (with morphological adaptations
for easier attachment to animal hair) more frequently form
part of the pollen spectra from the deeper parts of caves. This
fact may be caused by transport of pollen grains by animals.
In the Ramesh Cave (about 2000 m above sea level, sediments
dated 64–32 ka), Draxler (1992) interpreted the existence
of pollen from climatically demanding plants among other
types to be a consequence of the cave bear nourishment
(honey). The decrease in Pinus pollen recorded there is in
clear agreement with our results from Za Hájovnou Cave (see
above, Text-figs 2a, b). An overestimation of Cichorioideae
was visible only in layer 3ba (Text-fig. 1). Asteroideae
together with Pinus prevailed in most of the studied samples
from Za Hájovnou Cave.
According to Carrión et al. (1999) the pollen spectra from
cave sediments reflect the surface environment reliably only
when several requirements are fulfilled: a) taxonomic
diversity reliably above 15 taxons per sample, b) pollen
counts of more than 200 grains excluding Asteraceae, c) less
than 20% indeterminable pollen.
From cave sediments from the Moravian karstic areas not
only is the prevailing pollen from the Asteroideae but also
a more or less monotonous oryctocoenoses with overrepresentation
of smooth monolete Polypodiaceae spores,
Tilia and Corylus are also known (Doláková 2000, 2002,
2004, 2005). These pollen types are small and compact and
are known to be both mechanically and chemically resistant
(Havinga 1971, Jankovská 1971, Draxler 1992).
According to this information , the most reliable pollen
spectra which reflected the vegetation cover outside the caves
came from layers 2a and 2b (ZH P-8b and ZH P-5), the debris
cone, layer 4 (ZH P-10), and layer 5: 9-10m (ZH P-8b and
9). Care must be taken in interpretation of the vegetation and
climatic character of other studied palynospectra. In some
other site even a single common pollen grain could provide
useful information.
The wide variety and selectivity of the palynological
record from Za Hájovnou Cave did not allow accurate
reconstruction of changes in vegetation. The general
character of the pollen pictures corresponds to the climatic
optimum of the Holsteinian interglacial when compared to
studies of several localities in Central Europe (eg.
Dyjakowska 1952, Vodičková-Kneblová 1961, Břízová 1994,
Lang 1994, Kondratienė and Šeirienė 2003, Urban et al.
2011, Bittmann 2012, Urban and Sierralta 2012). The
findings of Pterocarya (often taken as a marker for the
Holsteinian) in the upper parts of the profile observed after
deterioration of the climate as reflected in the section of layer
5: 6.5–9 m from ZH P-9 (Text-fig.1) also support this
interpretation.
The samples from layer 5: 9.5–10 m of profile ZH P-9
and layer 4 lower of ZH P-8b support the Corylus expansion
described by Urban et Sierralta 2012 from Schöningen
lignite mine profile 12B LPAZ R 3b (MIS 9). The limited
palynological results from Za Hájovnou Cave do not provide
sufficient data to clearly differentiate if the sediments could
be related to MIS 11 or MIS 9; the assumption that they are
Holsteinien is a topical theme for discussion (eg. Geyh and
Müller 2005, Scourse 2006, Nitychoruk et al. 2006, Roe et
al. 2009, Bittmann 2012, Urban and Sierralta 2012).
The overall characteristics of the environment and
stratigraphic position based on other paleontological methods
is discussed in detail in a contribution by Musil et al. (2014).
According to faunistic and sedimentological characteristics,
the studied sediments can be divided into two groups.
Sediments older than the debris cone contained bones which
were nearly always complete, never weathered, and often
even in the correct anatomical positions. In sediments above
the debris cone the bones were invariably fragmentary,
many of them covered by sinter crusts. No fossils except
palynomorphs have been found inside the debris cone. The
difference between the palynospectra from the upper and
lower parts of the debris cone sediments could indicate
climatic variations during deposition of the debris cone (see
above, Text-fig. 1). Pollen spectra from lower layers and the
debris cone contained a limited number of Tertiary pollen
relics such as Pterocarya, Ilex, Celtis, which were not
observed in the older part of the sediments. These data are in
clear agreement with other Holsteinian localities (see above).
Conclusions
About 50 samples from profiles: ZH P-2, 5, 7, 8
and 9, 10 and 11 from Za Hájovnou Cave were assessed
palynologically.
The pollen spectrum of layer 7e is without any tree pollen
but with an accumulation of Asteroideae and several
hygrophilous herbs and thus may support the paleozoological
results suggesting development in a colder steppe
environment.
The general character of other palynospectra confirmed
the mild character of the climate during the Holsteinian
Interglacial due to the occurence of plants such as Carpinus,
Hedera, Tilia, Pterocarya and Ilex which are typical for the
climatic optimum of this time span. Such a pollen picture is
in clear accordance with other similar localities from Central
and Western Europe.
The difference in pollen record inside the debris cone
(layer 4) prompts division of layer 4 into 2 sublayers (layer
4 upper and layer 4 lower) which developed during varying
climatic conditions.
Detailed reconstruction of vegetations cover, their
changes and development is difficult to interpret from the
cave sediments. Selection, degradation and secondary
accumulation of various palynomorphs, due to their different
resistances to chemical and mechanical processes and
microbial attack during transport, were recorded. This
phenomenon was documented by numerical changes in the
different pollen grains from several spots within a single
layer in a direction towards the inner cave parts (primarily
a decrease in Pinaceae, and increase in Asteroideae).
38
The overall characteristics of the environment and
stratigraphic position based on other paleontological methods
will be discussed in the contribution Musil et al. (2014)
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40
Explanations of the plate
PLATE 1
Typical pollen grains (all magnifications 1 000 x)
1. Hedera sp. - layer 2b, profile ZH P-8b.
2. Pterocarya sp. - layer 2b, profile ZH P-8b.
3., 4. Tilia sp. - layer 5–9.5 m, profile ZH P-9.
5., 6. Carpinus sp. - layer 5–9.5 m, profile ZH P-9.
7. Juglans sp. – layer 4, profile ZH P-11.
8. Celtis sp. - layer 5–9.5 m, profile ZH P-9.
9. Quercus sp. - layer 4, profile ZH P-10.
10. Salix sp. - layer 5–10 m, profile ZH P-9.
11., 12. Corylus sp. - layer 5–9.5 m, profile ZH P-9.
13. Galium sp. - layer 3ba, profile ZH P-11.
14. Asteraceae Cichorioideae - layer 3b, profile ZH P-11.
15. Asteraceae Asteroideae - layer 2b, profile ZH P-8b.
16. Chrysosplenium/Ranunculus trichophyllum type layer
5–8.5 m, profile ZH P-9.
17. Poaceae - layer 5–9.5 m, profile ZH P-9.
18. Alnus sp. - layer 5–9.5 m, profile ZH P-9.
19. Pinus sylvestris type - layer 4, profile ZH P-11.
41
PLATE 1
42
Acta Palaeobotanica 47(1): 275–279, 2007
Use of fluorescent microscopy in the study
of redeposited palynomorphs in the cave and
marine sediments of Moravia (Czech Republic)
NELA DOLÁKOVÁ1
and ALENA BUREŠOVÁ2
1,2
Institute of Geological Sciences, Masaryk University, Kotlářská 2, 611 37 Brno, Czech Republic;
e-mail: nela@sci.muni.cz
Received 23 Jube 2006; accepted for publication 13 February 2007
ABSTRACT. Reworked palynomorphs occur in the cave sediments of the Moravian karstic areas as well as in
marine sediments of the Carpathian Foredeep (Czech Republic). Their preservation states may be different or
similar under the light microscope. The mutual distinguishing features of primarily Quaternary and Tertiary
ones, or of the palynomorphs from particular Miocene stages are therefore often very difficult to determine.
Observation under the fluorescent microscope can help to determine the reworked palynomorphs in Quaternary
as well as Miocene sediments.
KEY WORDS: fluorescent microscopy, redeposited palynomorphs, Miocene, Quaternary, Czech Republic
INTRODUCTION
It is not necessary for palynological studies
to be done only in autochthonous sediments
(i.e. of organic and chemical development).
The sediments develop mostly through disintegration,
transport and resedimentation of
older rocks. Therefore we can often find the
older redeposited palynomorphs in younger
sediments.
The occurrence of reworked palynomorphs
is typical for cave sediments. These sediments
do not contain plant remains in their original
positions. Palynomorphs are transported to the
sedimentation places with sedimentary particles
or through the activity of animals. These
facts create the possibility of mixing components
of different ages as well as selection of
palynomorphs due to their varied degrees of
resistance (Doláková & Nehyba 1999, Doláková
2000, 2002). The preservation state of the
redeposited grains can be similar to the grains
that are contemporary to the development of
the sediments and therefore it is frequently
difficult to mutually distinguish them.
Very similar problems arise in the Miocene
marine sediments from the Carpathian Foredeep.
Several transgression and regression
cycles occurred in this region. The redepositions
of foraminifers and calcareous nannoplankton
from the older Miocene stages into
the younger ones are commonly known from
this area (Brzobohatý et al. 2003). Thus, the
occurrence of redeposited palynomorphs is
likewise possible. The decision about whether
some pollen and spores typical for the climatic
zonations are in situ or not is therefore of great
importance. It is even important to diagnose
the potential amounts of redepositions within
frequent usual species, because the high percentage
of such redeposition could change the
image of palaeovegetation.
Observation under fluorescence microscopy
introduces a possibility to detect the reworked
palynomorphs. These methods were elaborated
mostly by van Gijzel (1967a, b, 1971,
1975, 1978).
All our macerated sediments were rich in
clays; they were partly calcareous, and for the
most part not coalified. For the maceration,
276
HCl, HF (not heating) and heavy liquid ZnCl2
were used. Pure glycerine was mostly used as
the observation medium. Part of the samples
(especially from the cave sediments) were
microscopically studied directly in ZnCl2 due
to the exclusion of further dilution of mostly
very small palynomorph amounts.
RESULTS
The sediments of the karstic formations
from the Moravian part of the Czech Republic
(Moravian, Javoříčko and Hranice Karsts)
are of Holocene, Pleistocene and Miocene ages
(Doláková 2000, 2002, 2004a, b, Doláková
& Nehyba 1999). The mixing of components
of different ages – especially Quaternary and
redeposited Tertiary – is common in these
areas. Observed under the light microscope,
their preservation states can be very similar.
It is therefore often very difficult to distinguish
the ages of individual palynomorphs
known both from the Quaternary and Tertiary
e.g. Pinus, Ulmus, Alnus, Quercus, Corylus,
and Betula, which consequently causes complications
in age determination and climatic
reconstructions. For example, 300 Quaternary
and 80 verifiable Tertiary palynomorphs were
determined in one sample from Ochoz cave
(Doláková & Nehyba 1999), their preservation
states being very alike (Pl. 1, fig. 3a, b).
Similar problems arise in the Miocene
marine sediments from the Carpathian Foredeep.
The occurrence of redeposited Cretaceous
palynomorphs is quite usual and well
detected. Under the optical microscope, the
existence of palynomorphs redeposited from
the older Miocene stages is often not possible
to prove. We try to use the observations under
the fluorescent microscope to detect some redeposited
palynomorphs (Burešová 2005).
Van Gijzel (1967a, b, 1971, 1975, 1978)
focused his attention on the methods of determining
the properties of UV-fluorescence on
fresh and fossil pollen and spores. He found
that the fluorescence spectra are closely
related to the chemical composition of palynomorphs.
They also depend on the different
levels of resistance to geological age, corrosion
and coalification of pollen and spore walls.
Similarly, weathering redepositions connected
with the oxidization of rocks and activities of
bacteria and fungi also change the intensity
and colours of the studied pollen and spores
(van Gijzel 1971). These characteristics are
then significant for determining the systematics
and age of palynomorphs. Most of these
methods require a lot of complicated measuring
and equipment (van Gijzel 1967a, b,
1975) for routine palynological studies. For
the orientational detection of the reworked
palynomorphs, the relative observation of the
colour spectra seem to be convenient. It is
necessary to attentively observe the change in
colour for a single type of palynomorph. With
increasing age, coalification and corrosion, the
colours shift from blue-green, white or yellow
and strong fluorescence to orange, red or
brown and weak fluorescence. Thus, each type
of secondary pollen shows a larger variation
in fluorescent colours than the autochthonous
material (van Gijzel 1971). New data about the
effect on fluorescence of the physical processes
associated with peat erosion and re-sedimentation
in reservoirs during the Holocene were
provided by Yeloff and Hunt (2005).
According to van Gijzel (1967b), the maceration
methods may also cause variation in
fluorescence (the use of hydrofluoric acid can
shift fluorescent colours of pollen to the red
end of the spectrum and darker). From our
experience, glycerine gelatine was found not to
be a suitable mounting medium for the study
of fluorescence. In a short time, the gelatine
changes colour (develops a dark circle) and the
palynomorphs fade. Glycerine as well as ZnCl2
seem to be suitable. The studied objects have
sufficiently stable colours for the duration of
observation.
The autochthonous pollen grains (except
grasses), both from the studied Quaternary
cave (Pl.1, fig.1a, b, 3a, b, 5a, b) and Miocene
marine sediments (Pl. 1, fig 2a, b), have light
colours (white, light-yellow, light-orange) and
intensive fluorescence during UV observation.
The pollen grains in the brackish more
coalified material from the Lower Miocene
sediments manifest shifts of colour towards
brownish-orange and a decreasing intensity of
fluorescence (Pl. 1, fig.6).
Verifiably redeposited palynomorphs from
the Moravian Quaternary karstic sediments
have typical dark brown colours with a very
low intensity of fluorescence (i.e. pollen grains
of Taxodiaceae – Pl. 1, fig. 3a, b). It is therefore
possible to assume the grains of similar
colours to also be redeposited (Pl. 1, fig. 1a, b).
277
Good identification of redeposition is possible
(from the different colours) in the mixture of
specimens of several ages of the one genus e.g.
Pinus from the Quaternary (Pl. 1) and from
the Miocene (Pl. 1, fig. 2) palynospectra. The
detection of single unusual pollen grains and
spores brought out more difficulties. Some
types of palynomorphs have primarily low
fluorescence or their exines are easier disintegrated
than others (i.e. Poaceae, Lygodium);
therefore, the redepositions are very hard to
confirm or eliminate.
Other interesting observations are connected
with the different colours of the algal
bodies. After Brooks (1971) and Yeloff and
Hunt (2005) the sporopollenin content of
lower plants is chemically different from that
of higher plants. The rate of corrosion of the
algal remains and fungal spores is different
from that of spores and pollen grains (van
Gijzel 1971); therefore, they differ considerably
in fluorescence from the other plant remains in
the fossil material. In the studied Quaternary
sediments (cave sediments, soils), the cenobia
of Botryococcus and Pediastrum show striking
blue-green colours (Pl. 1, figs 7, 8a, b). In the
Miocene sediments these were yellowish-white
with a very high intensity (Pl. 1, fig. 6). These
phenomena enable the detection of even small
parts of algae among other plant remains.
ACKNOWLEDGEMENT
We would like to express our thanks to Dr. P. Pokorný
and Dr. S. Sarkar (University Brno) for their
remittance to issues of fluorescence microscopy. This
study was supported by Grant Agency of the Czech
Republic, Grant 205/04/1021.
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z Ochozské jeskyně. (Sedimentological and Palynological
value of the sediments from the Ochozská
jeskyně cave). Geol.Výzk. Mor. Slez. in 1998: 7–10.
Brno. (in Czech).
Van GIJZEL P. 1967a. Palynology and flourescence
microscopy. Rev. Palaeobot. Palynol., 2:49–79.
Van GIJZEL P. 1967b. Autofluorescence of fossil pollen
and spores with special reference to age determination
and coalification. Leidse Geol. Meded.,
40: 263–317.
Van GIJZEL P. 1971. Review of the UV-fluorescence
microphotometry of fresh and fossil exines and
exosporia: 659–685. In: Brooks J., Grant P.R., Muir
M., van Gijzel P. & Shaw G. (eds) Sporopollenin.
Academic Press, London – New York.
Van GIJZEL P. 1975. Polychromatic UV-fluorescence
microphotometry of fresh and fossil plant
substances, with special reference to the location
and identification of dispersed organic material in
rocks: 67–91. In: Alpern B. (ed.) Colloque International
Petrographie de la Matiere Organique des
Sediments, Paris.
Van GIJZEL P. 1978. Recent developments in the
application of quantitative fluorescence microscopy
in palynology and paleobotany. Ann. Mines
Belgique, 7–8: 836–851.
YELOFF D.E. & HUNT C.O. 2005. Fluorescence
microscopy of pollen and spores: a tool for investigating
environmental change. Rev. Palaeobot.
Palynol., 133: 203–219.
278
Plate 1
1a. Part of pollen spectrum of the Late Glacial cave sediments – light microscope – red circle Pinus sp. autochthonous,
green circle Pinus sp. redeposited, Ochoz cave
1b. The same under the fluorescent microscope (UV light)
2a. Part of pollen spectrum from the Badenian marine sediments – light microscope – green circle redeposited
Pinus sp. (the cubic caves in the grain are caused by crystallization of pyrite in the anoxic marine environment),
Židlochovice
2b. The same under the fluorescent microscope (UV light)
3a. Redeposited Taxodiaceae from the Late Glacial cave sediments – light microscope, Ochoz cave
3b. The same under the fluorescent microscope (UV light)
4. Taxodiaceae from the Badenian marine sediments, the fluorescent microscope (UV light), Židlochovice
5a. Helianthemum sp. from the Late Glacial cave sediments – light microscope, Ochoz cave
5b. The same under the fluorescent microscope (UV light)
6. Pinus and Botryococcus sp. from Eggenburgian brackish sediments, the fluorescent microscope (UV light),
Trboušany
7. Cenobium of Pediastrum sp., fluorescent microscope, late Pleistocene, Krumlov Forest
8a. A – cenobium of Pediastrum sp. and B – several parts of another Pediastrum, fluorescent microscope, late
Glacial cave sediments, Ochoz cave
8b. A, B part of the same under the light microscope
P L A T E
279
N. Doláková & A. Burešová
Acta Palaeobot. 47(1)
Plate 1
N. Jb. Geol. Paläont. Abh. 271/2 (2014), 169–201 Article
Stuttgart, February 2014
The Badenian parastratotype at Židlochovice from the perspective of
the multiproxy study
Nela Doláková, Katarína Holcová, Slavomír Nehyba, Šárka Hladilová, Rostislav Brzobohatý,
Kamil Zágoršek, Juraj Hrabovský, Michal Seko, and Torsten Utescher
With 6 figures
Abstract: Two shallow boreholes were drilled in 2010 into the Badenian (Langhian) parastratotype
at Židlochovice (Carpathian Foredeep, NN5 Zone, Czech Republic). Their profiles (26 m of sediment)
were studied comprehensively (sedimentology, palaeontology – calcareous nannoplankton, red
algae, palynology, Foraminifera, Bryozoa, Brachiopoda, Ostracoda, Mollusca, Teleostei and Elasmobranchii).
The sedimentary succesion was biostratigraphically correlated with the NN5 Zone (14.9-
13.9 Ma), namely with the initial time of the Middle Miocene Climatic Transition. Seven lithofacies
representing multiple alternations of mudstone, sandstone and limestone facies were recognised
within sedimentary succession. Fossils generally indicated a normal marine, warm to subtropical environment.
The generally shallowing trend from the bottom (epibathyal/circalittoral) to the top (shallow
infralittoral) of the sedimentary succession with repeated palaeobathymetric changes could be
recognized in both boreholes. A mainly subtropical character of terrestrial flora was recorded. Within
this framework, either warm wet conditions with seasonal increases, or cooler phases were observed.
The abrupt change from mudstone deposited in a calm palaeoenvironment of the upper bathyal/circalittoral
to the variegated deposits of shallow water represents the most significant event correlable
with the FO of Orbulina (approximately 14.5-14.6 Ma). The interval below the FO of Orbulina can
be characterized by mudstone facies and significantly stable conditions of deposition, high nutrient
input and a decrease of oxygen content at the bottom. Seasonal stratification of the water column is
probable. Within this interval, cooling and an increase of seasonality were recorded. Above the FO of
Orbulina, there is evidence of shallowing connected with a higher flow regime and higher sedimentation
rate. The alternation of thick redalgal limestone bodies (a stable shallow palaeoenvironment with
low terrigenous input and seagrass meadows) and variegated sandstone, mudstone and limestone
interbeds (in an unstable deeper environment) possibly reflects orbitally forced climatic cyclicity.
Key words: Badenian, parastratotype, multiproxy quantitative analysis, palaeoecology, climate.
1. Introduction
The locality of Židlochovice – the clay pit of an old
brickyard (coordinates: long 16°37‘ 30‘‘; lat. 49°02‘
34‘‘; Z = 219-240 m a.s.l.) – is located at the southern
part of the Carpathian Foredeep in Moravia
(Czech Republic), which represents a peripheral system
of Central Paratethys basins (Fig. 1A). This site
has provoked the interest of many palaeontologists for
a very long time due to its rich fossil assemblages of
foraminifera, bryozoans, ostracods, microflora (nannoplankton
and red algae), molluscs (bivalves, gastropods,
cephalopods), sea urchins, corals, and the fish
otoliths and teeth (i.e., Procházka 1893; cicha et. al.
1956; PaPP et al. 1978; Říha 1983; BrzoBohatý 1997;
Doláková et al. 2008; zágoršek 2010a). Outcropping
deposits were assigned as a regional faciostratotype
of the Lower Badenian by PaPP et al. (1978) corres-
pondingtotheLanghianinthestandardchronostratigraphy
(hohenegger et al., in press).
©2014 E. Schweizerbart’sche Verlagsbuchhandlung, Stuttgart, Germany www.schweizerbart.de
DOI: 10.1127/0077-7749/2014/0383 0077-7749/2014/0383 $ 8.25
E
170 N. Doláková et al.
Fig. 1. A – Location of Židlochovice boreholes. B – Correlation of Židlochovice boreholes with biostratigraphical events
and global sequences 1 ranges from graDstein et al. (2012); 2 – Mediterranean ranges from Distefano et al. (2008); aBDul
aziz et al. (2008); hüsing et al. (2010); 3 – ranges from the Carpathian Foredeep (šváBenická 2002, this work); 4 – strati-
graphicalrangeofŽIDLboreholes;5–discrepancyinsuccessionofbiostratigraphicalevents(FOofOrbulinasuturalisand
LOofHelicosphaerawaltrans)betweenMediterraneanareaandŽidlochovicesuccession.
The Badenian parastratotype at Židlochovice 171
The outcrop sediments were biostratigraphically
correlated with the NN5 nannofossil biozone and
Orbulina suturalis (foraminiferal zone) (PaPP et al.
1978; Fig. 1B).They primarily consist of unlaminated
glauconitic calcareous clays – “tegel” – with intercalations
of calcareous sands or sandstone and limestone
with a dominance of red algae and bryozoans. The
environment was interpreted as a shallow circalittoral
on the basis of Foraminifera and Mollusca (PaPP et al.
1978). However, the study of faciostratotype from the
1970s is insufficient for current detailed interpretation.
Therefore, the goal of the revision was to reevaluted
biostratigraphy and to reconstruct an ecosystem evolution
in the faciostratotype locality using the multiproxy
quantitative analysis by trying to achive a
high resolution oft he records. In the present day, the
claypit is abandoned and covered with slope debris.
Therefore, two shallow boreholes representing the
original parastratotype section were drilled in 2010:
the lower interval was documented as ŽIDL-1 (11.9 m,
230 m a.s.l.) and the upper as ŽIDL-2 (16.9 m, 246 m
a.s.l.). The drilled cores were studied in detail from
the perspective of their sedimentology, palynology,
calcareous nannoplankton, red algae, Foraminifera,
Mollusca, Brachiopoda, Ostracoda, Bryozoa and Teleostei
– otoliths, teeth, Elasmobranchii (teeth). Some of
detailed palaeobiological analyses have been already
published (seko et al. 2012 – Ostracoda; Pavézková et
al., in press – Brachiopoda, tomaštíková & zágoršek
2012) – Bryozoa).
2. Geological setting
The locality at Židlochovice is located in the Carpathian
Foredeep (CF) near the boundary of the eastern
slopes of the Bohemian Massif that dip below Neogene
sediments and Carpathian flysh nappes (Fig. 1A).
The Badenian deposits of the CF in Moravia (Czech
Republic) represent the final stage of the depositional
history of the outer peripheral basins in the NW of the
Central Paratethys (meulenkamP et al. 1996). At the
beginning of the Middle Miocene, extensive erosion
took place in the Western Carpathians. The Badenian
transgression in the Moravian part of the Carpathian
Foredeep could be correlated with global sea level
cycle TB 2.4 (harDenBol et al. 1998, 14.8-13.6 Ma)
though local cycles were strongly controlled by tectonics
(kováč et al. 2001) and associated with basin
subsidence and continuous mountain chain uplift during
the synrift stage of back arc basin development.
The basins were filled with clastics transported from
uplifted areas (kvaček et al. 2006).
The studied sedimentation area was profoundly
differentiated. In general scheme at the beginning of
transgression in the shallow or elevated places, sand,
gravel, biostromal and biohermal limestone or calcareous
sandstone with a dominance of red algae and
bryozoas (Doláková et al. 2008; zágoršek 2010a)
were deposited. By contrast, a hundred meter deep depression
was filled with unlaminated calcareous clays
– “tegel” (BrzoBohatý 1997; kováč 2000; kováč et
al.2007).
3. Methods
A lithofacies analysis was conducted according to tucker
(1995), Walker & James (1992) and nemec (2005). The
shape and roundness of the coarsest grain fraction (> 4mm –
sieve separation, 17 samples) were estimated visually under
a microscope using the PoWers (1982) method. Combined
sieving and laser methods were used for grain size analysis
(22 samples). The Retch AS 200 sieving machine analysed
the coarser grain fractions (4 mm-0.063 mm, wet sieving);
the Cilas 1064 laser diffraction granulometer was used for
analyses of the finer fractions (0.0004-0.5 mm). Ultrasonic
dispersion, distilled water and washing in sodium polyphosphate
were used prior to the analyses in order to avoid
flocculation of the analysed particles. Average grain size is
expressed as the graphic mean (Mz), uniformity of the grain
size distribution/sorting as the standard deviation (σI) (folk
& WarD 1957).
Fossils (with the exception of palynomorphs, red algae
and calcareous nannoplankton) were studied from washing
residue (fractions 63-2000 µm) – 26 foraminifera samples
from the ŽIDL-l borehole and 24 samples from the ŽIDL-
2 borehole (Appendices 1, 2), molluscs (13+8) (Appendix
4), ostracods (13+8), brachipods (13+8), bryozoans (13+8),
bonefish and sharks (6+7) (Appendix 5), and fragments of
coralline algae. Calcareous nannoplankton was studied in
slides (26+24) (Appendix 3), coralline red algae in thin sections
(3+3) (Appendix 6), and palynomorphs after standard
palynological maceration in slides with pure glycerine observation
medium (16+16) (Appendix 7).
A KAPPA STM 723 stereomicroscope, Arsenal SZP
1102 ZOOM (bryozoans), WILD Heerbrugg (otoliths,
teeths), Nikon SMZ1 (molluscs, brachiopods), and Nikon
Alphaphot (red algae, palynomorphs) have been use for
taxonomic analyses.
The palynological problematic taxa identification was
done using the scanning electron microscope JEOL JSM –
649 OLV (Institute of Geological Sciences, Masaryk University);
ostracods – JEOL JSM 6390 (AVSR Banská Bystrica);
foraminifera – JEOL JSM 6380 LV (Charles University
in Prague); and bryozoan – low-vacuum SEM Hitachi
S3700N (Paleontological Department of the National Museum
in Prague). About 200300 specimens of foraminifera
from each sample were determined and the relative abundance
of taxa were calculated (Appendices 1, 2). The taphonomical
analysis of foraminiferal assemblages included the
study of abrasion and corrosion and the size sorting of tests
(holcová 1996, 1999). The palaeoecological interpretations
172 N. Doláková et al.
Fig. 2. List of lithofacies recognised in the drill holes Židlochovice 1 and 2 (Mz – mean/average grain size, σIstandart deviation/sorting
– counted according to folk & WarD (1957).
were based on the actuoecological data of culver & Buzas
(1980, 1981), Den Dulk et al. (1998, 2000), Jorissen et al.
(1995), kaiho (1994, 1997), van hinsBergen et al. (2005),
and murray (2006).
Slides for the study of calcareous nannoplankton were
prepared using the following technique: approximately 0.5
cm3
of rock sample was pulverized and watered with 5 ml
of water. One minute after shaking, one drop of suspension
from the middle of the water column was dripped on
a microscope slide. After this drying, standard microscope
slides were prepared and analysed using a light microscope
(normal and crossed nicols, 1000 x magnification). The
abundance of nannoplankton was expressed semiquantitatively
as a number of specimens in the visual field of the
microscope. The following categories were distinguished:
(i) very rare: 1-2 specimens; (ii) rare: 3-5 specimens; (iii)
common: 6-10 specimens; (iv) abundant: 11-20 specimens;
(v) very abundant: 21-40 specimens; (vi) mass occurrence:
above 40 specimens. The relative abundances of calcareous
nannoplankton species in assemblages were evaluated
quantitatively based on 200-500 determined specimens
from individual samples (Appendix 3).
Calcareous nannoplankton and foraminiferal assemblages
were statistically classified using the multivariate
techniques of PAST software (hammer et al. 2001). Euclidean
distance was chosen for quantification of object distance.
Four tested methods (PCA; cluster analysis: paired group,
single linkage, Wards method; nonmetric MDS, CABFAC
Factor analysis) gave comparable results. Finally, results of
the CABFAC Factor analyses were selected for publication,
because its results were the most illustrative.
Molluscs were evaluated semiquantitatively from 100 g
dry weight of sediment: less than 5 fragments were classed
as rare (1), 615 fragments as common (2), 1635 fragments as
abundant (3). Numbers of molluscan fragments reflect the
abundances of molluscan clasts among other bioclasts and
rock pieces in the washed residuum and do not express the
numbers of animals (Appendix 4).
A few samples of bryozoans from more lithified rock
were “laboratory weathered” and/or treated with acetic acid
as described by zágoršek et al. (2011). The method might
selectively dissolve some part of the skeleton unabling the
precise determination. However, without the treatment, the
number of determinable specimens considerable decreased.
Ostracod assemblages were analyzed with a focus on
taxonomy and palaeoecology, the later based on distribution
of the specimens and species along the borehole profiles, the
quantification of the valves/carapaces ratio, and the species
richness by using Simpson’s Reciprocal Index (seko et al.
2012).
For genera description of Rhodophyta, the methods by
Woelkerling (1988) and Braga et al. (1993) were used.
Microfacies were described according to flügel (2004).
PlanimetricanalysiswascalculatedwithJMicrovisionsoftware(roDuit
2008).
Standard maceration with HCl (20 %), HF, KOH and
The Badenian parastratotype at Židlochovice 173
HCl (10 %) and ZnCl2 (density = 2 g/cm3
) was used for the
palynological samples. Pure glycerine was most frequently
used as the observation medium. The percentage of individual
taxa was calculated from the total sum of a minimum
of 150 determined pollen grains and spores by using the
POLPAL programme (Walanus & nalePka 1999). The
palaeotropical and arctotertiary elements were classified
according to stuchlik et al. (1994). The terminology used
for vegetation units and partly for creating the pollen diagrams
follows kvaček et al. (2006) and kovar-eDer et al.
(2008). The pollen diagram was arranged into two parts due
to the overrepresentation of conifers: in the left section is
the pollen sum (100 %), excluding conifers, thus giving a
better visualisation of the basic character of the vegetation
changes. The right section shows the proportion of Pinus
and Cathaya and was counted from the sum of all grains.
4. Results
4.1. Sedimentarygeology
The lithofacies within the studied core profiles were
defined according to their grain size, rare preserved
sedimentary structures and petrology. Seven lithofacies
were recognised within the drill holes. Their characteristics
are presented in Fig. 2 and their distribution
can be followed in the presented lithostratigraphic
logs in Fig. 6A, B. Mudstone lithofacies (M1 and M2)
dominate in both drill holes, with a slightly higher in
drill hole ŽIDL-1, forming 69.7 % of the succession,
compared to 66.7% in drill hole ŽIDL-2.
The relative proportion of other lithofacies significantly
differs between the studied drill holes. The
ŽIDL-1 drill hole is characterized by a low content of
sandy facies (only 8.3 % of the profile and only S2
facies was recognized), a comparably higher content
of limestone facies L (17.9 %), and presence of heterolithic
facies (4.1 %). The ŽIDL-2 drill hole shows
a comparably higher presence of sandy facies (S1, S2,
S3; 22.0 % of the profile), lower content of facies L
(11.4 %) and the absence of the heterolithic facies.
Interpretation: The highly abundant mudstone lithofacies
reveal dominant deposition from suspension in
relatively calm conditions. Variations in the content of
sand, bioturbation, preservation of planar bedding and
shell debris also reflect periods of a relatively higher
input of material transported in traction (possibly by
storm currents). Sandy lithofacies and their alternation
with mudstone ones can be connected with deposition
in the lower shoreface or a transitional zone to a deeper
environment. The sharp bases of the beds, occurrence
of transported limestone clasts, shell debris and planar
lamination all support the role of an increased water
energy. The absence of clear wavy structures could
point to deposition below the normal wave base; we
can speculate about a high energy coast possibly non
barred. Heterolitic facies reflects the rapid alternation
of clastic input into the depositional environment.
The thick beds of limestone facies can be connected
with the stable conditions of deposition and also with
a severe reduction of clastic input. Thin limestone interbeds
could also be connected with the erosion and
redeposition of limestone into deeper environments
(possibly from the impact of storms). The multiple
alternation of relatively thin beds composed of mudstone,
sandstone and limestone facies (especially in the
ŽIDL-2 drill hole) are interpreted as cyclic changes of
depositional conditions (possibly climatically driven).
Repeated coarsening upward cycles in the successions
with the transition from mudstone facies to sandstone
and/or finally limestone can be interpreted as parasequences.
The significantly stable conditions of deposition
are presumed for the lower parts of the succession
in the ŽIDL-1 drill hole.
4.2. Biostratigraphy
Four planktonic foraminiferal and calcareous nanoplankton
bioevents were used for biostratigraphical
correlation (Fig. 1B)
(1) Praeorbulina circularis occurs from the base to
the top of both sedimentary successions
(2) The FO (i.e. First Occurrence) of Orbulina suturalis
was recorded at the level 5.7-5.8 m of the ŽIDL-1
borehole, in the ŽIDL-2 borehole, Orbulina suturalis
occurs from the base
(3) Neither Globorotalia preamenardii nor other
younger index species were found.
(4) Sphenolithus heteromorphus was recorded along
the whole sedimentary succession;
(5) Helicosphaera ampliaperta as well as Helicosphaera
waltrans, commonly and continuously occuring
in underlying deposits, were not recorded
(6) Uvigerina macrocarinata, the marker of the
regional Central Paratethyan substage (Moravian,
cicha et al. 1998), occurs from the base to the top
of both sedimentary succesion
(7) Among molluscs, a characteristic Badenian taxon
is Costellamussiopecten spinulosus.
The FO of the species “Chlamys” trilirata (ŽIDL-2)
lies above the base of the Middle Miocene in Paratethys,
and this species occurs only in the Lower and
?Middle Badenian (manDic 2004). Some of the pec-
174 N. Doláková et al.
Fig. 3. CABFAC Factor analysis (PAST softaware) of assemblages of benthic foraminifera, planktonic foraminifera and
calcareous nannoplankton assemblages.
tinid species at Židlochovice also occur in the Grund
Formation in the AlpineCarpathian Foredeep in
Lower Austria (correlated with the Lower Lagenidae
Zone – manDic 2004). Only Costellamussiopecten cf.
spinulosus – typical exclusively for the upper parts of
the Lower Badenian (Upper Lagenidae Zone – man-
The Badenian parastratotype at Židlochovice 175
Dic 2004) – is entirely absent in the Grund Formation,
whichindicatesthatthesediments atŽidlochoviceare
slightly younger than the sediments of the Grund Formation
(the situation is identical in Kralice nad Oslavou
– zágoršek et al. 2009).
Based on events (1) – (5), the ŽIDL-1 and 2 boreholes
can be correlated with the upper part of the NN5
Zone of calcareous nannoplankton (martini 1971)
above the LO of Helicosphaera waltrans and the upper
part of the M5 and lower part of the M6 zones
of planktonic foraminifera (Berggren et al. 1995).
According to the regional Central Paratethys chronostratigraphy
(using also regional bioevents (6) and
(7), the age of the sediments at Židlochovice can be
interpreted as Badenian, more exactly as upper part of
the Early Badenian (sensu PaPP et al. 1978; kováč et
al. 2007) or upper part of Mid Badenian according to
to the latest subdivision of the Badenian (hohenegger
et al. in press.).
4.3. Palaeobiology
4.3.1. Foraminifera and calcareous nannoplankton
Foraminiferal and calcareous nannoplankton assemblages
(Appendices 1, 2) were statistically classified
and results of CABFAC Factor analysis are presented
in Fig. 3:
Assemblages of benthic foraminifera can be well characterized
by four factors which explained 83.64 % of
variance; the 5th
and next factors explain each only less
than 1.5 % of variance.
(B1) The “Epiphytic” Factor 1 dominated by Asterigerinata
planorbis (4065%) may have alternated with
Elphidium spp. – Amphistegina spp. Factor 4. Epiphytic
species, which substantially dominated, allow to interpret
a well-aerated environment of seagrass meadows.
The assemblages often contain robust corroded
and abraded tests of shallow water species (mainly
Elphidium spp.) as well as foraminifera from different
palaeoenvironments (e.g. Uvigerina spp., Lenticulina
spp.) which indicates the postmortal transport of tests
by bedload in a higherenergy environment and/or reworking
of tests.
(B2) The “Highnutrient” Factor 2 with 30-55 % of
Pullenia bulloides, Nonion commune – Hansenisca
soldanii, Melonis pompiloides, Heterolepa dutemplei
and biserial textulariids: these high nutrient markers
(deep infauna, detrivore) indicate a high content of nutrients
in the sediment.
(B3) The “Cibicidoides – lowoxic markers” Factor 3 is
dominated by Bolivina dilatata, smallsized Cibicidoides
sp., Globocassidulina oblonga and Cassidulina
spp. (30-50 %). Foraminiferal tests are smallsized and
well preserved. The association of lowoxic infaunal
species along with opportunistic oxic species may indicate
(i) seasonal changes in water circulation – stratification
with hypoxic bottom environment change
during the year to nonstratified well oxygenated water
with a bloom of epifaunal suspension feeder, or (ii) hypoxia
only in the sediment (lowoxic infauna: Bolivina,
Globocassidulina and Cassidulina marker of phytodetritus
supply). At the bottom, there is a well aerated
environment with Cibicidoides sp. Generally, stress
conditions can be expected.
Statistical classification of planktonic foraminifera
by CABFAC Factor analysis is presented in Fig. 3.
Four factors explained 90-94 % of variance, % of variance
for the following factors does not exceed 2 %.
(P1) The smallsized Globigerina Factors 1 and 2
(Fourchambered G. praebulloides Factor 1 and G.
tarchanensis – Turborotalita quinqueloba Factor 2)
differs from the others in that its smallsized specimens.
Relative abundances of fourchambered globigerinas
reached values 15-35 % in samples with high
factor loading of the Factor 1 while fivechambered
globigerinids represent 50-80 % of assemblages with
high factor loading of the Factor 2. The relative abundancesofgroupsoffiveand
fourchamberedglobigeri-
nasnegativelycorrelated(correlationcoefficient0.52).
The both groups represent opportunistic, stresstolerant
assemblages, isotopic studies (holcová & Demeny
2012)indicatethe bloomof this smallsized specimens
to be probable after freshwater input. Fivechambered
globigerinids can indicate cold nonstratified water
(ruPP & hohenegger 2008); while Distefano et al.
(2010) described these foraminifers under hypersaline
conditions. Summarizing these interpretations, stress
condition represented mainly by salinity oscillations
can be expected for samples with high factor loadings
of these factors. Dominance of fivechambered globige-
rinidsmaybeaccompaniedwithmixedwatercolumn.
(P2) The Globigerina bulloides Factor 3 contains
common largesized Globigerina bulloides and Globigerinella
regularis (15-30 %) and characterized foraminiferal
assemblages generally with low P/Bratio
(to 20 %) and higher relative abundances of Globigerinoides
spp. and orbulinas (5-15 %). The warmwater
and oligotrophic species (Globigerinella regularis,
176 N. Doláková et al.
Globigerinoides spp. and orbulinas) (reynolDs &
thunell 1985; hemleBen et al. 1989; PuJol & vergnauD
grazzini 1995; sPezzaferri 1995; Bicchi et al.
2003; ruPP & hohenegger 2008) co-occurred with
eutrophic marker Globigerina bulloides (reynolDs
& thunell 1985; hemleBen et al. 1989) what may indicate
the seasonal succession of different groups of
plankton.
(P3) In samples with high factor loading of the Paragloborotalia
mayeri Factor 4, 25-60 % of nominative
species were recorded which can be accompanied by
small-sized Globigerina ex gr. praebulloides. The cooccurrence
of more eutrophic Globigerina spp. with
rather oligotrophic and warmer water Paragloborotalia
may indicate the succession of seasonal popula-
tions.
Calcareous nannoplankton assemblages can be excellently
classified by 3 factors which represent 99-75 %
of variance (Fig. 3):
(N1)ReticulofenestraminutaFactor1groupedassemblages
with more than 80% of small sized Reticulofenestra
minuta The small reticulofenestras are considered
to be stress-tolerant taxa indicating stress characterized
by quick changes within that environment,
including the oscillation of salinity (WaDe & BroWn
2006) and nutrient content (hallock 1987; Beaufort
& auBry 1992; flores et al. 1997; Wells & okaDa
1997;Bollmannetal.1998;kameo2002).
(N2) The Reticulofenestra haqii – R. minuta Factor 2
(small to medium-sized Reticulofenestra) is characterized
besides small R. minuta also by 5-15 % of specimens
from R. haqii group (sensu holcová 2012).
(N3) The Coccolithus pelagicus – Reticulofenestra
spp.Factor3 – is characterized byrelativeabundances
ofCoccolithuspelagicusover15%(15-45%)Thespecies
is a traditional indicator of cold and nutrient-rich
water (okaDa & mcinyre 1979; Winter et al. 1994),
but its common occurrence has also been recorded in
waters of up to 18 °C, which can be used as a tracer
of the periphery of areas of enhanced productivity
(cachao & moita 2000).
4.3.2. Mollusca
The systematic and palaeoecological evaluation of
molluscs is based on work by BagDasaryan et al.
(1966), PaPP et al. (1978), stuDencka (1986), stuDencka
et al. (1998), schultz (2001), manDic & harzhauser
(2003),andmanDic (2004).
Molluscs were present in all the studied samples of
both boreholes. The mollusc fauna consist predominantly
of bivalves; gastropods are less frequent. Small
gastropods of the genera Bittium, Alvania, Solariorbis,
Gibbula, or Rissoina were ascertained in some intervals
of both boreholes: ŽIDL-1: 2.7-2.8 m, 5.5-5.6 m,
8.4-8.5 m, 10.1-10.2 m; ŽIDL-2: 8.1-8.2 m, 8.7-8.8 m,
and 9.7-9.8 m.
The changing numbers of molluscan shells or their
fragments in the individual samples generally reflect
the changes in molluscan fauna abundance depending
on the changing palaeoecological conditions, mostly
the changes in water aeration.
In the basal part of ŽIDL-1, a low molluscan abundance
was ascertained (up to 7.2-7.3 m). In the upper
part, the amount of shells/fragments relatively increased
(maximum in 3.7- 3.8 m). In the rest of the profile
the amount of material varied. In the lower intervals
of the ŽIDL-2 profile, there was a low amount of
material (minimum in 15.9-16 m); in the upper part of
the profile, a gradual increase (maximum in 10.8-10.9
m) was observed, followed by a subsequent decrease
(the minimum at 8.7-8.8m).
Being an efficient active swimmer, Costellamussiopecten
prefers deeper calm waters without strong
currents and rather soft clay substrate (BagDasaryan
et al. 1966). “Chlamys” trilirata and Aequipecten
macrotis belong to epibionts usually exhibiting byssal
attachment to the substrate and needing primary
and secondary hard substrates for shell attachment.
They predominantly occur in rocky sublittoral environments
(manDic & harzhauser 2003), namely
in less exposed, deeper infralittoral (shallow subtidal)
zones. Cubitostrea digitalina is typical for the highly
exposed rocky medio/sublittoral (intertidal to shallow
subtidal) to depths of 10 m (manDic & harzhauser
2003). At Židlochovice, the oysters occurred more or
less continuously, but mostly in fragments. Nuculana
and Corbula represent sediment or suspension feeders,
shallowlyburrowing into muddy bottoms ofintertidal/subtidal
to bathyal depths. Nuculana is adapted
to stagnant waters with lower oxygen and higher hydrogen
sulphide contents. Corbula is generally an
opportunistic genus, optimally adapted to unstable
conditions.The predominanceofthin-shelledindeter-
minablebivalvefragmentsandsmallherbivoregastropods
in most samples indicates a calmer sedimentary
environment. The general presence of stenohaline bivalves
and occasional brachiopods, together with the
The Badenian parastratotype at Židlochovice 177
almost total absence of brackish and estuary elements,
confirms a fully marine (~35 ‰) sedimentary environ-
ment.
Among the bivalves, suspension feeders dominate,
confirming the environment to be rich in organic
detritus and planktonic microorganisms. The small
gastropods are mostly herbivores living in/on the algal
vegetation on the bottom (seagrass meadows). The
intervals with their presence can be – more or less –
correlated with a general decrease of shells/fragment
amounts, which can be interpreted as the consequence
of probable temporary lower water aeration (decrease
of O2 /increase of H2S content – stagnant water, ?greater
depth).
4.3.3. Ostracoda
Fiftytwo ostracod taxa were identified from the ŽIDL-
1 and ŽIDL-2 boreholes. The occurrence of “Gen.
indet.” moravica (Heliocythere moravica – paper in
preparation) confirms the early Badenian age of the
deposits (PaPP et al. 1978; seko et al. 2012). The sedimentation
rate affects the postmortal disintegration of
the carapace, and the increase of carapaces (valves/
carapace ratio) toward the upper part of both boreholes
might reflect an increase of the sedimentation
rate (oertli 1972). In the lower half of the ŽIDL-1
borehole, the amount of valves (33 to 301) is substantially
higher than the amount of preserved carapaces
(0 to 4). This ratio changes rapidly at a depth of 5.5-5.6
m, at which the carapaces once prevailed over valves
(25:49). In the remaining samples (toward the top),
an increase is still visible and remains stable with an
average value of 35 % of the carapace (valve:carapace
ratio) sample. A growing increase of carapaces (toward
the top) is also observed in the ŽIDL-2 borehole.
The maximum increase (39:104) occurs in the upper
part 8-18.2 m (seko et al. 2012).
Another remarkable change was observed at the
depth of 5.5-5.6 m of the ŽIDL-1 borehole. A turnover
in the assemblage composition indicates a transition
between two marine habitats (seko et al. 2012). In the
lower part of the ŽIDL-1 borehole (12-7.2 m), we observed
a higher amount of specimens like Buntonia
subulata subulata, Cytherella pestiensis postdenticulata,
Henryhowella asperrima, Bosquetina carinella,
Parakrithe dactylomorpha and Krithe sp. which according
to gross (2006) are indicative of deeper circalittoral
to epibathyal environments. On the contrary,
the occurrence of epineritic Cytheridea acuminata
(12-7.2 m; not only) could indicate a transition to an
infraneritic environment, supported by the presence of
epineritic to infraneritic Aurila opaca, Cnestocythere
lamelicosta. There was a significant increase in specimens
of Aurila species, Cnestocythere lamellicosta,
Loxocorniculum hastatum, Pokornyella deformis,
Senesia philippi and Tenedocythere sulcatopunctata,
which prefer epineritic environments and tolerate a
decrease in salinity (gross 2006). The disappearance
of circalittoral to epibathyal ostracods indicates the
transition to shallow infrallitoral (5.6-0.9 m). In the assemblage
of the ŽIDL-2 borehole, a decrease in the
abundance of deeper circalittoral to epibathyal species
(Henryhowella asperrima) is observed toward the top.
Furthermore, indicators of a shallower palaeoenvironment
(Senesia philippi) and infralittoral and phytal
species (Cnestocythete lamellicosta) became dominant
part of assemblage. In comparison to the ŽIDL-1
borehole, the change in species dominance and composition
is not so significant. Aurila species dominate
in the samples and in this parameter the assemblage
of the ŽIDL-2 borehole is similar to the assemblage
of the upper part of the ŽIDL-1 borehole (5.6-0.9 m),
reflecting shallow infralittoral conditions (seko et al.
2012).
The ostracod assemblages have been analysed
according PSH Platycopids Signal Hypothesis (Whatley
et al. 2003). The percentage of Platycopida is
not higher then 11.5 %, indicating a well oxygenated
palaeoenvironment (oxygen content ›5 ml/l; Whatley
et al. 2003). This interpretation does not agree with
foraminifera and mollusc assemblages which indicate
a poorly oxygenated environment. This contradiction
can be explained either by results of modern data,
which unsupported PSH (BranDão & horne 2009)
or by seasonal variations of the oxygen content at the
seafloor and in the sediment.
4.3.4. Bryozoa
tomaštíková & zágoršek (2012) identified 116 bryozoan
taxa; 23 of them are mentioned for the first time
in this locality. Bryozoans dominate in all the studied
samples: cheilostomes represent almost 70 % (81 species),
while only 35 species of cyclostomes were found.
Among the determined bryozoan species, those from
temperate and tropical environments dominate. Very
common are Metrarabdotos maleckii, Steginoporella
cucullata, S. tuberculata and Adeonella polystomella,
characteristic for recent tropical seas (moissette
1988).
Generally, the diverse bryozoan fauna indicate normal
marine, temperate to subtropical environments.
Based on bryozoan growth form analysis (according to
178 N. Doláková et al.
hageman et al. 1997 and mckinney & Jackson 1989),
shallow water prevails. The most common bryozoan
colonial growth forms are erect; encrusting growth
forms are less abundant. There were only two reteporiforms
and no free living growth forms were found.
The absolute number of species of entire growth form
depends on the preservation (zágoršek et al. 2012a).
However, the oscillation in the ratio between erect
and encrusting forms in the same sedimentary succession
may indicate changes in the environment. In the
lower part of the ŽIDL-1 borehole, palaeoecological
analyses based on the evaluation of the ratio between
erect and encrusting bryozoans indicate lower temperatures
at the sea bottom because of greater water depth
(80-100 m), thus resulting in a large amount of erect
bryozoans. In higher parts of the ŽIDL-1 and ŽIDL-2
profiles, minor differences in the ratio of both growth
types confirm a relatively stable, rather shallow and
warm sea environment with estimated depth about 20-
30 m (as similarly interpreted also in Přemyslovice see
zágoršek et al. 2012b).
The lower part of the ŽIDL-2 borehole (a depth
of 17-15.7 m) shows a remarkably large proportion
of erect bryozoan growth forms, the largest being at
a depth of 16 m (19 erect and only 4 encrusting species
were identified). The increase in the proportion of
erect growth forms may indicate an increase of depth
and therefore a decrease of temperature at the sea
floor, which is comparable to the interval of the ŽIDL-
1 borehole at the depth of 9.3 m.
In the following sequences, the ratio between the
erect and encrusting bryozoan growth forms gets closer:
at the depth of 12.8 m it is almost the same (26/24),
showing what is probably the shallowest environment
and therefore the warmest water in the studied sedimentary
succesion, comparable to those at the depth
of 3.8 m in the ŽIDL-1 borehole.
In samples from depths of 7.3 m in ŽIDL-1 and 10.9
m in ŽIDL-2, the total number of species rapidly decreased.
This may indicate an unsuitable environment;
probably the increase of water energy also corresponds
to a higher amount of reteporiforms bryozoans in these
samples. The second event is represented by samples
at 3.8 m in ŽIDL-1 and 9.8 m in ŽIDL-2, where the
diversity of the bryozoan community is the highest.
These samples may be interpreted as bryozoan events
comparable with those described by zágoršek (2010b)
from other sections of the Carpathian Foredeep.
Summarizing, the lowest part of the studied sedimentary
succesion (the lower part of the ŽIDL-1
borehole culminating at 9.3 m) represents a deeper
and therefore cooler environment similar to those
described for example from Korytnica (zágoršek et
al. 2012a) than the rest of the profile. Conversely, the
younger part (the upper part of the ŽIDL-2 borehole
culminating at 12.8 m) indicates the shallowest and
warmest environment similar to those reported from
Přemyslovice (zágoršek et al. 2012a) in comparison
with the rest of the profile.
4.3.5. Brachiopoda
Brachiopods have been studied for the first time (Pavézková
et al., in press) at Židlochovice. There were 4
speciesidentified,allfromthefamilyMegathyrididae,
namelyArgyrothecacuneata,Argyrothecasp.,Joania
sp., and Megathiris detruncata. In both boreholes,
brachiopods occur in negligible amounts, although
Argyrotheca cuneata is relatively the most numerous
species. All species recognized in Židlochovice
are generally common in the Miocene sediments
of the Central Paratethys. Extant representatives of
Megathyrididae are mostly shallow water species,
exhibiting a cryptic mode of life (logan 1979). The
occurrence of brachiopods at Židlochovice – in spite
of their scarcity – significantly supplements not only
total spectrum of Badenian fauna from this locality,
but also mosaic of Badenian brachiopod occurrences
in the Carpathian Foredeep on the territory of the
Czech Republic.
4.3.6. Ichthyofauna
Ichthyofauna was represented intermittently by otoliths
(14 species) and singularly by the isolated teeth
(1 species) of teleosts and sharks (1 species) in both
boreholes. Due to poor taxonomic representation in
the small size of samples from the boreholes, the results
are approximate.
The ŽIDL-1 borehole shows the occurrence of
otoliths of mesopelagic fishes such as Valenciennellus
tripunctulatus, Diaphus acutirostrum, D. cahuzaci,
D. regani and Notoscopelus mediterraneus
throughout the section accompanied by teeth of the
bathypelagic shark Deania sp. at 2.8-3.0 m. Such an
assemblage is indicative of the deepest sublittoral with
good communication with mesopelagic environments
(BrzoBohaty 1997). Picture deviation from this trend
only occurs at 10.6-10.8 m, where shallow water taxa
such as Lesueurigobius ex gr. vicinalis and maybe
even a juvenile specimen of Acropomatidae indet. are
present, possibly indicating an episodic shallowing of
the environment. This also seems to be in accordance
The Badenian parastratotype at Židlochovice 179
Fig. 4. Co-occurrence of foraminiferal and calcareous nannoplankton clusters (Fig. 3) and their schematized position in
the basin.
with the numerous gypsum fragments in the washed
residuum from this level.
Mesopelagic fish otoliths persist in the lower part
of the ŽIDL-2 borehole, where epimesopelagic Vinciguerria
poweriae, Benthosema fitchi and Diaphus
div. sp. dominate. At the same time, some sublittoral
elements (Brachydeuterus, Gobiidae) occur. The upper
part is characterised by sublittoral elements – teeth
of sparids, otoliths of juvenile gobiids and mere fragments
of juvenile mesopelagic diaphids.
Taking into account the original palaeobathymetric
analyses based on otoliths from the Židlochovice
brickyard, the deepest environment in the bottom
part of the original loam pit (150-250 m, BrzoBohatý
1997) was clearly indicated by adult otoliths of the
nonmigrating bathydemersal macrourid Trachyrincus
scabrus. To the top of the whole Židlochovice section,
the fish fauna reflects a general shallowing from
the shallow bathyal/deeper circa-littoral to the shallow
sublittoral. The ichthyofauna in the whole section indicates
normal salinity.
4.3.7. Red algae and microfacies
Red-algal limestone consists of coralline algae (12.5-
57.7 %) and bryozoans (1.9-38.3 %). Other determined
components (foraminifers, echinoids, polychaetes,
molluscs), if present at all, represent less than 3 %.
However, the volume of unrecognized fragments in
some samples is 0.5-51 %. Lithoclasts can constitute
0.18-2 %. Micrite is always present in 2.25-3 % and
sparite forms in 0.14-8 %. The limestone is of low porosity
(0.2-5 %).
Samples can be classified as coralline algal and
coralline algal – bryozoans microfacies. Composition
of samples with respect to the identified facies is given
in Appendix 6. The growth forms of the coralline
algae are monospecific protuberant rhodoliths, multispecific
rhodoliths with the nucleus of fragments of
other frequently protuberant corallines, simple abraded
protuberances and the debris of them. Rhodoliths
are mostly found in fine biogenic sandy matrices rich
in micrite. On the contrary, protuberances are usually
180 N. Doláková et al.
Fig. 5. Pollen diagram (A) arranged after paleoecological groups: left side – pollen sum (100%) excluding conifers, the right
side proportion of Pinus and Cathaya (100% all grains).
Zidlochovice CA palaeoclimate data (B): Temperatures: MAT- mean annual temperature, CMT- temperature of the coldest
month, WMT- temperature of the warmest month. Precipitation: MAP- mean annual precipitation, thick line segments: all
taxa, thin line segment: wet taxa excluded.
accompanied by lithoclasts.
Although coralline algal assemblages consist of six
genera: Lithothamnion, Mesophyllum, Phymatolithon,
Lithophyllum, Spongites and Sporolithon, the first two
mentioned dominate in the assemblage. It can be concluded
that the algae of the subfamily Melobesioideae
The Badenian parastratotype at Židlochovice 181
predominateoverthesubfamiliesofLithophylloideae,
Mastophoroideae and Sporolithoideae in all samples.
This assemblage is frequent in nontropical marine environments
(aguirre et al. 2000). Although coralline
algaespeciesdescriptionsare beyondthe scopeofthis
paper, it should be noted that Phymatolithon calcareum
(Pallas) aDey & mckiBBin was observed by as
monospecificabraded protuberances as wellas crusts
in multispecific rhodoliths.
According to the presented data, three limestone
facies can be evaluated. First, rhodolith floatstone
(RFgm) in a grainstone matrix consists of multispecific
and monospecific rhodoliths floating in a fine-grained
biogenic sandy matrix (ŽIDL-1, 6.3 m). Lithoclasts are
present as low amounts values or are absent. These facies
indicate deposition in the outer shelf environment
with evidence of storms events (flügel 2004).
Second, coralline algal detritus limestone (CAdl)
consists of abraded protuberances; however, few
multispecific rhodoliths are present (ŽIDL-1: 4.5-5
m, ŽIDL-2: 10.6 m and 13.3 m). Facies correspond
to coralline algal grainstone and to lithoclastic coral-
linealgallimestone.Othercomponentsarelithoclasts,
unsorted allochems, sparite, and micrite. The facies
indicate an environment of middle to inner shelf, with
higherterrigenousinput(flügel 2004).P.calcareum
indicatesa depth of 36-93m(Basso 1998).
Third, bioclastic breccias (Bbr) consist of angular
fragments of rhodoliths and molluscs (ŽIDL-2, 9.6 m).
The matrix is mainly micritic, without lithoclasts or
mediumsized allochems.
The limestone facies successions thus indicate
changes of depositional environment in the drill cores.
The transition from rhodolith floatstone to coralline
algal detritus limestone from the ŽIDL-1 interval of
6-34.5 m indicates the shallowing of the environment.
4.3.8. Palynology
The most diversified pollen spectra have been found
in environments with reduced oxygen content (ŽIDL-
1: 11.9-7.5 m and ŽIDL-2: 16.81-5.6 m), also evident
as cubic caves where there were pyrite crystals in the
pollen grains. The overdominance of conifers and
frequent dinoflagellates with a very small amount of
other pollen and spores were observed in ŽIDL-1 at 76
m and in ŽIDL-2 at 15.26 m. The remaining samples
were sterile or very poor in palynomorphs.
A mainly subtropical character of flora was interpreted
(Fig. 5). Three zonal forest formations were
spread around the basin. Subtropical Broadleaved
forests comprised up to 30 % of the evergreen elements
(100 % without Pinus, Cathaya): including
Sapotaceae, Engelhardia, Platycarya, evergreen
Fagaceae – Castanopsis, Trigonobalanopsis, morphotypes
Tricolporopollenites henrici, T. microhenrici,
Tricolporopollenites liblarensis, Reevesia, Cornus/
Mastixia, Rutaceae and Araliaceae, Pteridaceae (Fig.
5A). Proportions of most thermophilous elements of
P1 sensu stuchlik et al. (1994) were slightly lower in
comparison with the Lower Miocene palynospectra
of the CF (Doláková et al. 1999, 2011; Doláková &
slamková 2003; kováčová et al. 2011).The share of deciduous
woody elements resulting in warm temperate
Mixed Mesophytic and Broadleaved Deciduous forest
types (i.e. Quercus, Carya, Celtis, Juglans, Tilia,
Betula, Acer) was lower in all the studied samples (to
a maximum of 17 %). A higher diversity of “oak type”
pollen grains (e.g., Quercus robur/pubecscens) has
been recorded in comparison with the Lower Miocene
sediments from CF (Doláková et al. 1999; kováčová
et al. 2011).
The fluctuation of coastal swamp (Taxodiaceae,
Cyrillaceae, Myricaceae) and riparian elements (Ulmus,
Fraxinus, Liquidambar, less generous Alnus, Salix,
Lythraceae) could be as a result of humidity changes
(our interpretation). Pollen grains of Platanus represent
a marked component of the pollen spectra. Herbs and
heliophilous elements such as Poaceae, Asteraceae,
Caryophyllaceae, Chenopodiaceae and Ericaceae are
consistently present. Urtica, Plantago, Thalictrum,
Salvia, Lavandula and Ephedra have been recorded
locally. The sporadic occurrence of Botryococcus and
the pollen of the aquatic coastal plants Sparganium,
Potamogeton and Nymphaea indicate freshwater inflow
in some facies.
Frequent various Pinaceae (Pinus, Cathaya) were
components of mountain coniferrich forest (with the
admixture of Cedrus, Tsuga, and Picea in higher altitudes)
and along with Sciadopitys they were also a part
of the intrazonal lowland formations.
The development of the main climatic characteristic
is expressed in the graphs of the Coexistence
Approach (Fig. 5B). The general climatic conditions
are in accordance with Bruch et al. (2004). Warm
wet conditions can be observed in the lowest interval
of the studied sequence (ŽIDL-1: 11.9-10.6 m). The
seasonality increases in the middle part of ŽIDL-1
(with well expressed seasonality of precipitation –
the driest month ~2040 mm; the wettest month 200-
250 mm); this is followed by a cooler phase evident
from the upper part of ŽIDL-1. A slightly increasing
portion of arctotertiary elements is also visible from
182 N. Doláková et al.
Fig. 6. A, B – Correlation of bioevents an lithology within studied groups of fossils. a) ŽIDL-1. b) ŽIDL 2. Cadl: coralline
algal detrital limestone, Carg: coralline algal branch rudstone to floadstone and grainstone, RF: rhodolith floatstone in grainstone
biodetrital matrix, E: epiphytic gastropods.
the palynograph (Fig. 5A). The repeating of warm wet
conditions is recorded in the lowest part of ŽIDL-2:
16.8-15.7 m.
5. Discussion
5.1. Age
Thedatingofglobalbiostratigraphicaleventsrecorded
in the studied succession differs in the world’s oceans
(graDstein etal.2012)fromthatoftheMediterranean
area (aBDul aziz et al. 2008; Distefano et al. 2008;
hüsing et al. 2010). The differences are summarized
inFig.1B.Due to the communication of the Central
Paratethyan Sea with the Mediterranean, the correlation
of bioevent timing with the Mediterranean dates
is rather probable. The succession of the guide bioevents
– the LO of Helicosphaera ampliaperta which
precedes the FO of Orbulina suturalis – agrees with
the Mediterranean area. However, co-occurrence of
the Helicosphaera waltrans and Orbulina suturalis
described from the Mediterranean was not recorded in
the faciostratotype at Židlochovice. This discrepancy
caused that lower boundary of studied succession can
be dated only approximately: (i) in case of later appearance
of Orbulina suturalis in the study area, the
age of this boundary must be younger than 14.36 Ma
(= the LO of Helicosphaera waltrans); (ii) more probably
age ranges between 14.91 Ma (= the LO of Heli-
The Badenian parastratotype at Židlochovice 183
cosphaera ampliaperta) and c. 14.5-14.6 Ma (= FO of
Orbulina suturalis in Mediterranean area). The upper
boundary is older than the FO of Globorotalia praemenardii
which is dated in the Mediterranean area to
13.92 Ma (Distefano et al. 2008).
5.2. Climate
The changing of warm wet conditions, seasonality in-
creasesandacoolerphasewithinthemainsubtropical
character of terrestrial flora could represent the final
phase of the Miocene Climatic Optimum Böhme
(2003), utescher et al. (2000), Bruch et al. (2010).
Decreasing of the most thermophilous elements in
comparisonwiththe LowerMiocenepalynospectraof
the CF, higher diversity of “oak type” pollen grains
and evidence of the NS oriented climatically dependent
gradient(Doláková et al. 1999;Doláková et al.
2008;kováčová etal.2011;Doláková &kováčová,in
prep.) indicate the beginning of the Middle Miocene
Climatic Transition (14.8-12.0 Ma) (floWer & kennett
1994, see also harzhauser et al. 2011).
The overdominance of conifers was similarly
found in some other pollen spectra from CF from the
NN5 zone (e.g. Oslavany, Rebešovice). From palaeoecological
point of view, this can be caused for several
reasons: the morphology of the land (sandy dunes or
mountainous areas), huge pollen production and the
great amount of air transport. Accumulation in marine
sediments distant from the seashore and greater resistance
to oxygenation could also play a role (heusser
1978, hoPkins & mccarthy 2002). This interpretation
is supported by the synchronously plentiful occurrence
of dinoflagellates.
184 N. Doláková et al.
5.3. Marine environment
The co-occurrence of calcareous nannoplankton and
foraminiferal assemblages distinguished by factor
analysis in the samples made it possible to compile the
model of marine environment (Fig. 4):
(1) The highnutrient benthic foraminiferal assemblages
accompanied by the calcareaous nannoplankton assemblages
with higher abundances of Coccolithus pelagicus
represent the deepest water assemblages recorded
only in mudstone. As Coccolithus pelagicus is a marker
of eutrophic waters, it agrees with the expected high
amount of nutrients in the sediment; The “Cibicidoides
– lowoxic marker” Factor cooccurs mainly with the
planktonic assemblages dominated by small fivechambered
globigerinids. Both assemblages tolerate unstable,
stress conditions (oscillations of salinity and/or nutrient
supply). Cooccurrence with Turborotalita quinqueloba
may indicate mixed water column.
(2) The epiphytic foraminifera represent the shallowest
environment with seagrass meadows recorded
mainly with the Reticulofenestra minuta and with all
planktonic foraminifera assemblages. Because epiphytic
foraminifers indicate shallow water sea (depth
in the first tens of meters, murray 2006), the postmortem
transport of planktonic foraminifera by superficial
currents and/or the reworking of plankton is expected,
which agrees with the variegated planktonic assemblages.
Horizons with smallsized foraminiferal tests
within the limestone body (4-6 m of ŽIDL-1 borehole)
indicate a sizesorting of tests due to transport in suspensions
(i.e. storms) or in flows. The epiphytic foraminifera
occur in limestone and sand.
The distribution of fossil assemblages along the studied
sedimentary successions (Fig. 6A, B shows the alternation
of three main types of palaeoenvironments:
(1) The interval below the FO of Orbulina (below 6.8
m of the ŽIDL-1 borehole) can be characterized by
mudstone lithofacies deposited from suspension in relatively
calm conditions. Rich benthic foraminiferal assemblages
with a higher abundance of infauna, a high
nutrient marker including elongate biserial textulariids
(mainly Spiroplectinella) indicate a rich source of nutrients
in the sediment; all these species are detrivores;
the majority of them can tolerate a decrease of oxygen
content in accord with the presence of Mollusca. The
higher abundance of Coccolithus pelagicus in the calcareous
nannoplankton assemblages also indicates a
higher nutrient source for primary producers. On the
other hand, the oligotrophic planktonic foraminifera
indicate episodical stratification. Episodic increase of
Spiroplectinella abundance (above 5 %) imply that this
event depend on specific palaeoecological conditions
(? supply of specific nutrients) and cannot be isochronous
biostratigraphical zone (Spiroplectammina Zone
sensu grill 1941). The predominance of Ostracoda
valves indicates a slow sedimentation rate. Ostracoda,
Bryozoa, molluscs and fish otoliths and teeth indicate
a deeper environment. There is a decrease of oxygen
caused by the absence of algae and other organic material
creating a suitable surface for encrusting bryozoans.
Ichthyofauna proved to have good communication
with the mesopelagic environment. Diversified terrestrial
vegetation with up to 30 % of broadleaved evergreen
and thermophylous elements, a lower proportion
of deciduous woody taxa of zonal angiosperms and
swamp, and riparian and heliophylous azonal taxons
were observed in the pollen spectra. The changing of
the warm wet conditions, seasonality increases, and
cooler phase were recognized. The deepest environment
in the bottom part of the original loam pit of
the Židlochovice brickyard (150-250 m, BrzoBohatý
1997) was documented.
At the levels of 9.8 m and 8.9 m, the shortterm deviation
from a fully marine environment are suggested
by elevated levels of stresstolerant foraminifera and
rare opportunistic mollusc assemblages. The postmortal
transport of foraminiferal tests is very probable,
which also confirms the sedimentological record with
variations in the content of sand, bioturbation, preservation
of planar bedding and shell debris, reflecting
periods of higher input of material transported by current
in traction (possibly during storms). The seasonality
increase followed by beginning of the cooler phase
were recorded in pollen spectra.
(2) The upper interval with Orbulina is characterized
by a variegated palaeoenvironment with rapid alternations
of lithofacies. Generally, two horizons can be
distinguished:
(2a) Thick beds of limestone can be connected with
stable conditions of deposition and with the reduction
of clastic input. Epiphytic foraminifera as well as
phytal ostracoda and epiphytic gastropods indicate the
occurrence of seagrass meadows. Abraded and corroded
tests reflect a high-energy environment around the
wave base and shallowing. Limestone with red algae
is present. Ostracoda, Bryozoa, molluscs and fish otoliths
and teeth indicate shallow water conditions. The
The Badenian parastratotype at Židlochovice 185
increased ratio of Ostracoda valves/carapaces indicates
the rapid burial of shells, or the general increase
in the depositional rate; this could also be caused by
rapid episodic sedimentation after intensive episodic
rainfalls.
Palynomorphs, when present, are marked by the
dominance of conifers and marine Dinoflagellata.
These accumulations (similar to some others of the
same age in the CF – Oslavany, Rebešovice) may be
caused for palaeoecologic or taphonomic reasons.
(2b) Sandy lithofacies and their alternation with mudstone
and thin limestone interbeds were deposited
in a higher flow regime. The absence of clear wavy
structures points to deposition below the normal wave
base. Heterolitic facies support the rapid alternation
of clastic input in the depositional environment. Thin
limestone interbeds could also be connected with erosion
and the redeposition of limestone into a deeper
environment (possibly as a result of storms). These
lithofacies mainly contain opportunistic small-sized
Cibicidoides and the lowoxic markers indicate the instability
of the environment, the coexistence of oxiphylic
taxa at the seafloor, and lowoxic conditions in
the sediment, or seasonal changes of oxygen content
at the seafloor due to the seasonal stratification of the
water column. Common Globocassidulina and Cassidulina
is considered to be a marker of phytodetritus
input, which may be seasonal. Palynospectra are mostly
impoverished (such as in 2a). Only at the interval of
15.6-16.8 m from ŽIDL-2 there were well diversified
pollen and spores indicating warm wet conditions.
6. Conclusions
1) Two shallow boreholes recording 26 m of sediment
were drilled in 2010 on the Badenian parastratotype
at Židlochovice (Carpathian Foredeep, NN5 Zone,
Czech Republic). In comparison with the original definition
of the parastratotype:
– The sedimentary succession was studied using sedimentological
and palaeontological methods.
– All systematic groups were newly reworked (calcareous
nannoplankton, Foraminifera, Ostracoda, Mollusca,
Teleostei except Anthozoa and Echinoidea), or
extended (red algae, Bryozoa, Brachiopoda, Elasmobranchii,
pollen grains); detailed quantitative study
was used.
– The environment of sedimentation and its development
over time was newly interpreted.
– The stratigraphical position of the parastratotype
was stated more precisely.
2) Biostratigraphical dating of the sedimentary succession
enabled a correlation with the NN5 Zone 14.9 to
13.9 Ma, namely with the initial time of the Middle
Miocene Climatic Transition.
3) Seven lithofacies were recognised within the sedimentary
succession representing multiple alternations
of mudstone, sandstone and limestone facies. The
fauna generally indicated a normal marine, warm to
subtropical environment. Individual groups of fauna
and flora confirm what appears to be a generally shallowing
trend from the bottom (epibathyal/circalittoral)
to the top (shallow infralittoral) of the sedimentary
succession with repeated palaeobathymetric changes
in both boreholes. Though influence of local tectonic
movements is expected, the general shallowing may
be correlated with regressive phase of the TB 2.4 cycle
dated on 14.2-13.6 Ma (harDenBol et al. 1988).
A mainly subtropical character of terrestrial flora
within warm wet conditions, seasonality increases and
a cooler phase were observed.
The most significant event correlable with the FO
of Orbulina (c. 14.5-14.6 Ma) is the abrupt change from
mudstone deposited in the calm palaeoenvironment of
upper bathyal/circalittoral to the variegated deposits of
shallow water.
The lower interval below the FO of Orbulina can
be characterized by mudstone facies and significantly
stable conditions of deposition, high nutrient input and
a decrease of oxygen content at the seafloor and/or in
sediment. Episodical stratification of the water column
is probable. Interval cooling and an increase of seasonality
were recorded.
Based on FO of Orbulina, shallowing connected
with a higher flow regime and a higher sedimentation
rate are supported. The alternation of thick redalgal
limestone bodies (a stable shallow palaeoenvironment
with low terrigenous input and seagrass meadows) and
variegated sandstone, mudstone and limestone interbeds
of an unstable deeper environment could reflect
orbitally-forced climatic cyclicity.
Acknowledgements
The research is supported bythe Grant Project 205/09/0103
(Grant Agencyof the Czech Republic)and MSM0021620855.
We are grateful to the native speaker lector Mrs. sarka
rousava for correction of the English language. milos Bartol,
Patrick grunert and mathias harzhauser are gratefully
acknowledged for valuable remarks and comments
in their reviews, and günter schWeigert for his help with
the editorial arrangement.
186 N. Doláková et al.
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Manuscript received: September 24th, 2013.
Revised version accepted by the Stuttgart editor: December
9th, 2013.
Addresses of the authors:
nela Doláková, slavomír nehyBa, rostislav BrzoBohatý,
JuraJ hraBovský, Institute ofGeological Sciences,Masaryk
University,Kotlářská 2,61137Brno,CzechRepublic;
e-maisl: nela@sci.muni.cz, slavek@sci.muni.cz
katarína holcová, Institute of Geology and Paleontology,
Charles University in Prague, Albertov 6.128 43 Praha 2,
Czech Republic;
e-mail: holcova@natur.cuni.cz
šárka hlaDilová, Department of Biology, Faculty of Education,
Palacky University, Purkrabská 2, 771 40 Olomouc,
Czech Republic;
e-mail: sarka.hladilova@upol.cz
kamil zágoršek, Department of Paleontology, National
Museum, Václavské nam. 68, CZ 115 79 Prague, Czech
Republic;
e-mail: kamil_zagorsek@nm.cz
michal seko, Geological Institute, Slovak Academy of
Sciences, Ďumbierska 1, 974 01 Banská Bystrica, Slovakia;
e-mail: michal.seko47@gmail.com
torsten utescher,SteinmannInstitute,UniversityofBonn,
Nussallee 8, Bonn, Germany;
e-mail: utescher@geo.unibonn.de
190 N. Doláková et al.
Appendices
Appendix 1: List and relative abundances of benthic foraminiferal species.
Z1/11.5-11.6m
Z1/10.2-
10.3m
Z1/9.8-
9.9m
Z1/9.0-
9.1m
Z1/8.9-
9.0m
Z1/8.5-
8.6m
Z1/7.8-
7.9m
Z1/7.2-
7.3m
Z1/6.8-
6.9m
Z1/6.3-
6.4m
Z1/6.3m
Z1/6.0-
6.1m
Z1/5.7-
5.8m
Ammoniabeccarii(linne) 0.38 0 1.16 0 0 0 0 0 0 0.45 0 0 0
Ammoniatepida(cushman) 0 0.88 0 0 0 0.81 0.4 0 0.45 0 0 0 0.66
Amphicoryna badenensis (D’orBigny) 0.38 0 0 0 0 0 0.4 0 1.35 5.41 0.76 0 0
Amphistegina bohdanowiczi BieDa 0 0 0 0.39 0 0 0 0 0 0 0 0 0
Amphisteginamammilla(fichtel etmoll) 0 0 0 0 0 0 0 0 0 0.9 0 0 0
Angulogerina angulosa (Williamson) 0 0 0 0 0 0 0 0 0 0 0 0 0
Asterigerinata planorbis (D’orBigny) 0 5.31 12.79 6.25 12.02 4.86 2.82 0.87 1.35 20.27 26.72 31.39 39.07
Baggina sp. 2.26 0 0.78 0.39 0 0.81 2.82 1.74 1.35 0.9 0 0 0
Bolivina antiqua D’orBigny 0 0 0 1.17 0 0.81 0 0.43 0 0.9 0.38 0.73 0
Bolivinadilatatadilatatareuss 0.75 8.41 17.44 1.56 11.24 0 2.42 1.3 3.59 0 0.38 0 0.66
Bolivina hebes macfaDyen 0 0 0 0 0 0 0 0 0 0 0.38 0 0
Bolivinaplicatellacushman 0.75 3.54 0.78 0.39 1.16 4.45 4.03 3.04 1.79 0 0 0 0
BolivinapokornyicichaetzaPletalova 0 0 0 0 0.39 0 0 0 0 0 0 0 0
Bolivinascalprataretiformiscushman 0 0 0 0 0 0 0 0 0 0 0 0 0
Buchnerinabuchneri(margerel) 0.38 0 2.33 0.39 2.33 0 0.4 0.43 1.79 0 0 0 0
Bulimina elongata D’orBigny 6.39 4.87 6.98 2.34 5.43 6.88 4.03 0.43 0 2.25 0.38 0.73 3.97
Bulimina striata D’orBigny 6.02 0 1.55 1.17 0.78 0.81 0.4 4.78 2.69 4.5 1.15 0 1.32
Cancris auriculus (fichtel et moll) 1.13 0 0 0 0.78 0 0.4 0 0 0 0.76 0 0
CassidulinalaevigataD’orBigny 0.75 6.19 2.71 2.73 2.33 1.62 2.02 3.48 1.35 1.35 0.38 0 4.64
Cibicides sp. (small-sized) 0 5.31 8.53 2.34 13.95 2.02 0.81 3.91 4.93 0.9 13.74 10.95 19.21
Cibicidoides austriacus (D’orBigny) 0 0 0 0 0 0 0 0 0 0 0 0 0
Cibicidoides ungerianus (D’orBigny) 0.38 0.88 0.39 0 1.55 0 0 0.87 1.35 6.31 7.25 10.22 3.97
Dentalina sp. 0 0 0 0 0 0.4 1.21 0 0.45 0.45 0 0 0
Elphidiumcrispum(linne) 0 0 0 0.39 0 0 0 0 0 0.9 0.38 0 0
Elphidium fichtellianum (D’orBigny) 0 0.88 0.39 0 0 0 0 0.43 0.45 1.8 1.15 0 0.66
Elphidium flexuosum (D’orBigny) 0 0 0 0 0 0 0 0 0 0 0 0 0
Elphidiummacellumfichtel etmoll 0 3.54 1.16 1.95 1.55 3.64 1.61 1.3 0 7.21 8.4 13.14 5.96
Elphidiumortenburgense(egger) 0 0 0 0 0 0 0 0 0 0 0.38 0 0
Elphidium rugosum (D’orBigny) 0 0 1.16 0 0 0.4 0.81 0 0 0 1.15 3.65 1.99
Elphidium sp. (juvenile) 0 0 0 0 0 0 0 0 0 0 0 0 3.97
Ehrenberginaserratareuss 0 0 0 0 0 0 0 0 0 0 0 0 0
Fursenkoina acuta (D’orBigny) 1.13 0.44 0 1.17 0 0.4 0.81 0.87 0 0 0 0 0
Chilostomellaovoideareuss 0 0 0 0 0 0 0 0 0 0 0 0 0
Globocassidulinaoblonga(reuss) 2.26 3.98 5.43 3.52 6.98 3.64 5.24 8.7 4.04 0.9 5.34 0.73 2.65
Globulina gibba D’orBigny 0 0 0.78 0 0 0 0 0 0.9 0.45 0 0 0
Globulina spinosa D’orBigny 0 0 0 0.39 0 0 0 0 0 0.45 0 0 0
Grigelis pyrula (D’orBigny) 0 0 0 0 0 0 0.4 0 0 0 0 0 0
Hansenisca soldanii (D’orBigny) 3.01 9.29 0.78 5.08 2.33 3.24 4.84 5.22 5.83 1.35 1.15 2.19 0
Hanzawaiaboueana(D’orBigny) 1.13 2.65 0.78 1.56 0.78 0 0.4 1.3 0.45 3.15 1.53 0.73 1.99
Heterolepa dutemplei (D’orBigny) 11.65 7.08 5.04 15.23 7.75 11.74 8.06 10.43 8.07 4.05 0.76 5.11 0
Heterostegina sp. 0 0 0 0 0 0 0 0 0 0 0 0 0
Hoeglundina elegans (D’orBigny) 0.75 0.88 0 0.78 0.78 0.4 3.63 1.74 1.35 1.35 0.76 0.73 0
Karreriellachilostoma(reuss) 0.38 0 0 0 0 0 0 0 0 0.9 0.38 0 0
Laevidentalina elegans (D’orBigny) 0 0 3.49 0.39 1.16 1.62 0 0 0 0 0 0 0
Lagena striata (D’orBigny) 0.38 0.44 0 0 0 0 0 0.43 0 0 0 0 0
Lagena hexagona (Williamson) 0 0 0 0 0 0 0 0 0 0 0 0 0
Lenticulinacalcar(linne) 0 0 0.39 1.95 0 2.83 0 0.87 3.14 0 0 0.73 0
Lenticulina clypeiformis (D’orBigny) 0 0 0.39 0 0 0 0 0 0 0 0 0 0
Lenticulina inornata (D’orBigny) 0.38 0 0.78 0.78 0.78 0 2.02 0.87 0.9 1.35 0 0 0
Lenticulina orbicularis (d´orBigny) 0 0 0 0 0 0 0 0 0 0 0 0 0
Lenticulina sp. (broken or abraded) 0 0 0 0 0 0 0 0 0 0 0 1.46 0
Lenticulina sp. (juvenile) 0 0 0 0 0 0 0 0.43 0 0 1.91 0 0
Lenticulina vortex (fichtel et moll) 0 0 0 0 0 0 0 0 0 0 0 0 0
Lobatula lobatula (Walker et JacoB) 0.75 1.33 0.39 1.95 1.16 3.64 0.81 0 0 14.41 11.45 10.22 7.28
Marginulina hirsuta D’orBigny 0 0 0 0 0 0 0 0 0 0 0.38 0 0
The Badenian parastratotype at Židlochovice 191
Appendix 1: continued
Z1/5.1-
5.2m
Z1/4.8-
5.0m
Z1/4.3-
4.4m
Z1/4.2-
4.3m
Z1/4.0-
4.2m
Z1/3.6-
3.7m
Z1/3.1-
3.2m
Z1/2.6-
2.8m
Z1/2.2-
2.3m
Z1/2.0-
2.1m
Z1/1.9-
2.0m
Z1/1.7-
1.8m
Z1/1.2-
1.3m
Z1/1.0-
1.1m
Paleoecological characteristic
(Jorissenetal.1992,1995,2007,
kaiho 1994, De stigter et al. 1998;
De Dulk et al. 2000, sPezzaferri et
al.2002,van hinsBergen etal.2005,
murray 2006, BálDi 2006)
0 0 0.93 0 0 0 0 0 0 0.37 0 0 1.23 0.39 Euryhaline
0 0 0 0 0 0 0 0 0 0 0 1.74 0 0.39 Euryhaline
0 0 0 0 0 0 0 0 0 0 0.63 0 0 0.39
51.28 5.26 12.09 0 0.49 0 2.14 1.82 36 5.9 7.59 1.74 2.87 4.31 Oxiphylic
0 0 0 0 0 0 0 0 0 0 0 0 0 0 Oxiphylic
0 0 0 0 0 0 2.5 0 0 0 0 0 0 0
22.22 52.63 46.05 12.39 37.93 14.29 14.64 15 0 40.96 29.75 21.3 23.77 13.73 Epiphytic
0 0 0 0 0 0 0 0 0 0 0 0.43 0 0
0 0 0 1.71 0 3.67 0.36 0.91 0 1.11 0 0 0 0.39 Hypoxic, infauna
0 0 0 6.84 0 11.02 7.86 11.36 0.5 0.37 0.63 8.26 1.64 1.57 Hypoxic, infauna
0 0 0 0 0 0 1.07 0 0 0 0 0 0.82 0.78
0 0 0 3.42 0 0 3.57 0.91 0.5 0 0 3.48 2.05 0.39
0 0 0 0.43 0 0 0.36 0 0 0 0 0 0 0 Hypoxic, infauna
0 0 0 0 0 0 0 5 0 0 0 0 0.41 0
0 0 0 0.43 0 1.22 1.07 2.27 0 0.37 0.63 2.61 2.46 0.39
0 0 0.93 0 0.99 3.67 2.5 3.18 0 1.11 0 3.91 2.46 0.78 Hypoxic, infauna, high-nutrient
0 0 0.47 3.42 0 0 6.79 3.64 0 0.74 2.53 1.3 1.64 3.92
0 0 0 0 0 0 0 0 0 0 0 0 0 0.39
0 0 0 10.26 0.49 4.49 6.79 4.09 0 0 0 6.09 4.92 4.31 Phytodetritus supply
0.85 9.47 4.19 10.26 12.81 9.8 4.29 10.91 1.5 2.95 0 11.74 15.98 7.45 Oxiphylic, stress-tolerant
0 0 0 0 0 0 0 0 0 0 0 0 0.41 0.39 Oxiphylic
0 5.26 4.19 4.7 6.4 4.9 3.57 6.36 6.5 1.85 6.96 7.83 5.74 2.35 Oxiphylic
0 0 0 0 0 0 0 0 0.5 0 1.27 0 0 0.78
10.26 11.58 15.35 1.71 10.84 8.57 3.57 1.82 8 7.38 6.33 2.17 4.1 2.75 Euryhaline
0 0 0 0.43 1.97 1.63 0 0.45 0.5 1.85 1.9 1.74 2.87 2.75 Euryhaline
0 0 0 0.43 0 0 0 0 0 0 0 0 0 0 Euryhaline
0 0 0 0 0 0 0 0 0 0 0 0 0 0 Euryhaline
0 0 0 0 0 0 0 0 0 0 0 0 0 0 Euryhaline
0 0 0.47 0 0 0.41 2.86 0 0 0 0.63 0.87 0.41 1.57 Euryhaline
0 0 0 0 0 0 0 0 0 0 0 0 0 0 Euryhaline
0 0 0 0 0 0 0 0 0 0 0 0 0 0
0 0 0 0.43 0 0 0 0 0 0 0 0 0 0 Hypoxic, infauna,
0 0 0 0 0 0 0 0 0 0 0 0 0 0 Hypoxic
0.85 0 0.47 11.97 0 5.71 3.57 4.55 0.5 3.32 1.27 2.61 1.64 2.35 Phytodetritus supply
2.56 0 0 0 0.49 0.41 0.36 0 0 0 0.63 0 0 0.39
0 1.05 1.86 0.43 0 0 0 0 2 0 0 0 0 0
0 0 0 0 0 0 0 0 0 0 0 0 0 0
0 0 0 1.71 0 1.63 2.14 0.91 1 0.37 0.63 0.43 0.41 3.14 Hypoxic, infauna, high-nutrient
0.85 2.11 0 0.43 0 1.22 2.5 0.45 0 1.48 0 0 1.64 1.57
5.98 5.26 4.65 2.56 1.48 2.04 0.36 2.27 12.5 4.43 8.23 2.17 2.87 2.75 Oxiphylic
0 0 0 0 0 0 0 0 0 0 0 0 0 0 Oxiphylic
0 0 0 0 0 0 1.07 0 0 0 0 0 0.82 3.53
0 0 0 0 0 0 0 0 0.5 0 0 0 0 1.18
0 0 0 0 0 0 0 0 0 0 0 0 0 0
0 0 0 0.43 0 0 0 0 0 0 0 0.43 0.41 0.39
0 0 0 0 0 0 0.36 0 0.5 0 0 0 0 0
0.85 0 0 0 0 0 0 0 0 0.74 0 0 0 1.18
0 0 0 0 0 0 0 0 0 0 0 0 0 0
0 0 0 0.43 0 0.82 0.36 0.45 1 0.37 0.63 0.43 0 0
0 0 0.47 0 0 0 0 0 0 0 0 0 0 0
0 0 0 0 0 0 0 0 0 0 0 0 0 0
0 0 0 0.43 0 0 0 0 0 0 0 0 0 0
0 0 0 0 0 0 0 0 0 0 0 0 0 0
4.27 5.26 5.12 7.69 20.69 18.37 8.93 5 8.5 11.81 6.33 5.65 6.56 7.45 Oxiphylic
0 0 0 0 0 0 0 0 0.5 0 0 0 0 0
192 N. Doláková et al.
Appendix 1: continued
Z1/11.5-11.6m
Z1/10.2-
10.3m
Z1/9.8-
9.9m
Z1/9.0-
9.1m
Z1/8.9-
9.0m
Z1/8.5-
8.6m
Z1/7.8-
7.9m
Z1/7.2-
7.3m
Z1/6.8-
6.9m
Z1/6.3-
6.4m
Z1/6.3m
Z1/6.0-
6.1m
Z1/5.7-
5.8m
Martinottiella karreri (cushman) 0 0 0 0 0 0 0 0 0 0.9 0 0 0
Melonis pompiloides (D’orBigny) 0.75 1.77 1.55 1.95 0.78 2.83 2.02 4.78 8.07 1.35 3.44 0 0
Nonion commune (D’orBigny) 10.15 3.1 5.04 3.52 4.65 7.29 7.26 6.52 7.17 1.8 2.29 0 0.66
Nonion sp. 0 0 0 0 0 0 0 0 0 0 0 0 0
Nodosaria hispida d´orBigny 0 0 0 0 0.78 0 0.4 0.43 0 0 0 0 0
Nummoloculinacontraria(D’orBigny) 0 0 0 0 0 0 0 0 0 0 0 0 0
Oridorsalisumbonatus(reuss) 0 0 0 0 0 0 0 0 0 0.45 0 0 0
Neugeborina longiscata (D’orBigny) 0 0 0 0 0 0 0 0 0 0 0 0 0
Pappina parkeri (karrer) 0 1.77 0 0.78 0 0.81 0.4 0.43 0 0 0 0 0
Pararotalia aculeata (D’orBigny) 0 0 0 0 0 0 0 0 0 0 0 0.73 0
Planularia lanceolata (D’orBigny) 0 0 0 0 0 0 0 0 0.45 0.45 0 0 0
Plectofrondicularia digitalis (neugeBoren) 2.26 0.88 0.78 2.73 0.78 1.21 0 0.87 2.24 1.35 0 0 0
Praeglobobuliminapyrula(D’orBigny) 1.88 1.33 0 0.39 1.55 3.64 1.21 1.74 0.9 0.9 0 0 0
Porosononion granosum (D’orBigny) 0 0.44 0 0 0.39 0 0.81 0.87 2.24 0 0 0.73 0
Pseudotriloculina consobrina (D’orBigny) 0 0.44 0 0.39 0 0 1.21 0 0.9 0.45 0.38 0 0.66
Pullenia bulloides (D’orBigny) 5.64 0.44 2.71 5.86 2.33 2.83 4.03 3.91 1.35 2.25 2.67 0.73 0
Pyramidulina raphanistrum (linne) 0.75 0 0 0.78 0 0 0 0.43 0 0 0 0 0
Quinqueloculina buchiana d’Orbigny 0 0 0 1.95 0 0 0 0 0.45 0 0 0 0
Quinqueloculina hauerina (d’Orbigny) 1.88 0.44 0 2.34 0.78 4.05 2.42 0.43 1.79 0 0 1.46 0
Reussellaspinulosa(reuss) 0 0 0 0.39 0 0 0 0 0 0 0.38 0 0
Rosalina sp. (cf. semiporata (egger) 0 0 0 0.39 0 0 0 0.43 0 0.45 0.38 1.46 0
Semivulvulina deperdita (D’orBigny) 11.28 0 0 0 1.94 3.64 6.45 6.09 4.48 0.45 1.15 0 0
Sigmovirgulina tortuosa (BraDy) 0 0 0 0 0 0 0 0 0 0 0 0 0
Siphonina reticulata (czJzek) 0 0 0 0 0 0 0 0 0 0 0 0 0
Sphaeroidina bulloides D’orBigny 0.38 0 0 0 0 0 0.4 0.87 0.45 1.8 0.38 0 0
Spiroloculina sp. 0 0.44 0 0.78 0.39 0.4 1.61 0 0 0.45 0.38 0 0
Spirorutilus carinatus (D’orBigny) 1.13 2.65 1.16 6.25 1.94 6.88 8.47 8.26 2.24 0 0 0 0
Stilostomella adolphina (D’orBigny) 2.63 6.19 3.1 4.69 1.94 2.02 2.82 2.61 2.69 0 0.38 0 0
Stilostomella elegans (D’orBigny) 4.14 0 3.1 1.17 2.71 3.24 3.23 0 2.24 1.35 0 0 0
Textularia gramen D’orBigny 0 0.44 1.55 1.56 1.16 1.62 4.03 1.74 0 0 0 0 0
Textularialaevigata D’orBigny 0.38 0.44 0 0 0 0 0 0 0.45 0 0 0 0
Trifarina bradyi cushman 0.75 0.44 1.55 0 0.39 0 0 0.43 0 1.8 0 0 0
UvigerinaaculeataD’orBigny 0 0.44 0.78 2.73 1.94 2.02 1.61 3.04 9.42 0.45 0 0 0
Uvigerinabononiensisfornasini 0 0 0 0 0 0 0 0 0 0 0 0 0
UvigerinamacrocarinataPaPP etturnovsky 14.29 0 0 3.13 0.39 2.43 0.81 2.17 4.93 0.45 0.76 0.73 0
Uvigerina pygmoides PaPP etturnovsky 0 0 0 0 0 0 0 0 0 0 0 0.73 0
Uvigerinasemiornatad´orBigny 0 0 0 0 0 0 0 0 0 0 0 0.73 0
Valvulineria complanata (D’orBigny) 0.38 12.39 2.33 3.91 0 0 0 0 0 0 0 0 0.66
P/B-ratio 13.64 41.6 32.28 20.74 37.38 40.05 29.34 48.2 44.25 43.22 30.13 9.87 11.7
Foraminiferal number
(specimens/1 g of rock sample)
308 348.3 571.5 129.2 374.55 529.71 1263.6 799.2 720 156.4 75 15.2 34.2
The Badenian parastratotype at Židlochovice 193
Appendix 1: continued
Z1/5.1-
5.2m
Z1/4.8-
5.0m
Z1/4.3-
4.4m
Z1/4.2-
4.3m
Z1/4.0-
4.2m
Z1/3.6-
3.7m
Z1/3.1-
3.2m
Z1/2.6-
2.8m
Z1/2.2-
2.3m
Z1/2.0-
2.1m
Z1/1.9-
2.0m
Z1/1.7-
1.8m
Z1/1.2-
1.3m
Z1/1.0-
1.1m
Paleoecological characteristic
(Jorissenetal.1992,1995,2007,
kaiho 1994, De stigter et al. 1998;
De Dulk et al. 2000, sPezzaferri et
al.2002,van hinsBergen etal.2005,
murray 2006, BálDi 2006)
0 0 0 0 0 0 0.36 0 2 0 1.9 0 0 0.39
0 0 0 3.85 0 0.41 1.07 1.82 3.5 0.37 7.59 0 1.23 5.49 Hypoxic, infauna, high-nutrient
0 0 0 0.43 0.49 1.63 0.71 1.36 0 0.37 0.63 0 0 1.57
0 0 0 0 0 0 0 0 0 0 0 0 0 0
0 0 0 0 0 0 0 0 0 0 0 0 0 1.18
0 0 0 0 0 0 0 0 0 0 0 0 0 0
0 0 0 0 0 0 0 0 0 0 0 0 0 0
0 0 0 0 0 0 0 0 0 0 0 0 0 1.57
0 0 0 0 0 0 0.36 0 0 0 0 0 0 0
0 0 0 0 0 0 0 0 0 0 0.63 0 0 0
0 0 0 0 0 0 0 0.45 0 1.11 0 0 0 0
0 0 0 0.43 0 0.41 0.36 0.91 0 1.11 0 0 0 0
0 0 0 0 0 0 0 0 0 0 0.63 0 0 0,39 Hypoxic, infauna, high-nutrient
0 0 0 0 0.99 0.41 1.79 3.64 0.5 0.74 0 1.74 0.41 0 Euryhaline
0 0 0.47 0 0 0 0 0 0.5 0 0 0 0.82 0
0 0 0 0 0 0.41 1.07 0.45 0 0.74 0.63 0.43 0 0 Hypoxic, infauna, high-nutrient
0 0 0 0 0 0 0 0 0 0 0.63 0 0 0
0 0 0 0 0 0 0 0 0.5 0.37 1.9 0 0 0,39 Oxiphylic
0 1.05 0 0 0 0 0 0 0 0 0 0 0 0 Oxiphylic
0 0 0 2.56 0 1.22 1.07 2.27 1.5 0.74 0 4.35 3.69 3,53
0 0 0 0 0 0 0 0 0.5 0.74 0 0 0 0
0 0 0 0.43 1.97 0 1.43 0 1.5 0.37 0 0 0 0
0 0 0 0 0 0 0 0 0 0 0 0 0.82 0
0 0 0 0 0 0 0 0 0 0 0 0 0 0
0 0 0 0 0 0 0 1.82 0.5 0.37 0 0 0.41 0,78
0 0 0 1.28 0 0 2.14 0.91 0 0.37 0.63 0.43 0.82 0,78
0 0 0 0 0 0 0.36 0.45 1 0 0 0.87 0 1,18
0 0 0 4.27 0.49 1.22 4.29 4.55 2 1.85 1.9 3.48 2.87 4,31
0 0 0 0.43 0 0 0.36 0 1 1.11 1.9 0 0 0
0 0 0 0 0 0 0 0 0 1.85 2.53 0.43 0.41 0
0 0 0 0 0 0 0 0 0 0 0 0 0 1,18
0 0 1.4 2.56 1.48 0.41 0 0 0 0 0 0 0 0,39
0 0 0 0 0 0 0 0 0 0 0 0 0 1,18 Hypoxic, infauna, high-nutrient
0 0 0 0 0 0 0 0 0 0 0 0 0 0 Hypoxic, infauna, high-nutrient
0 0 0 0.43 0 0 1.07 0 3 0.37 1.9 0.87 0 0,39 Hypoxic, infauna, high-nutrient
0 0 0.93 0 0 0 0 0 0 0 0 0 0 0 Hypoxic, infauna, high-nutrient
0 0 0 0 0 0 0 0 0.5 0 0 0 0 0 Hypoxic, infauna, high-nutrient
0 1.05 0 0.43 0 0 0 0 0.5 0 0 0.43 0.41 1,18 Hypoxic, infauna, high-nutrient
0 1.04 40.27 41.65 1.93 29.55 34.73 48.39 32.43 16.36 37.55 42.29 31.46 34,28
23.4 9.6 132.6 243.03 41.4 130.95 107.25 263.03 59.2 116.64 25.3 160.8 160.2 279,36
194 N. Doláková et al.
Z2/16.9-17.0m
Z2/16.6-16.4m
Z2/15.9-16.0m
Z2/15.4-15.5m
Z2/15.1-15.2m
Z2/14.4-14.8m
Z2/14.1-14.2m
Z2/13.9-14.0m
Z2/13.8-13.9m
Z2/13.7-13.8m
Z2/13.3-13.4m
Z2/12.9-13.0m
Z2/12.2-12.3m
Z2/11.5-11.6m
Z2/11.0-11.2m
Z2/10.8-10.9m
Z2/9.9-10.0m
Z2/9.2-9.3m
Z2/8.8-8.9m
Z2/8.5-8.6m
Z2/8.3-8.4m
Z2/7.8-7.9m
Z2/6.9-7.0m
Z2/6.0m
Ammoniabeccarii(linne) 0 0 0 0 0 0 0 0 0 0.42 0 0 0 0 0 0 0.75 0.82 0 0 0 0 0 0
Ammonia tepida (cushman) 1.12 2.09 2.87 0 1.51 0 0.41 0 0 0 0.4 0 0.81 1.32 0.98 0 0 0 0 0.8 0 0.42 0 0
Amphicoryna badenensis (D’orBigny) 0 0.42 0.41 0 0 0.42 0 0 0 0 0 0 0 0 0 0.45 0 0 0 0 0 0.84 0 0.37
Amphistegina bohdanowiczi BieDa 0.37 0 0 0 0 0 0.82 0 1.41 5.02 1.62 1.52 0 0 2.44 3.13 6.02 6.58 3.42 1.2 4.29 0.84 0 0
Amphisteginamammilla (fichtel etmoll) 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.41 0 0 0.48 0 0 0
Angulogerina angulosa (Williamson) 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1.12
Asterigerinata planorbis (D’orBigny) 12.3 11.3 13.5 38.9 32.1 47.1 7.76 6.43 14.1 11.3 32.4 14 4.03 4.82 42.4 31.3 22.9 36.2 31.2 29.7 38.6 26.4 1.35 0.75
Baggina sp. 0 0.84 0 0 0 0 0.41 0 0 0 0 0 0 0 0 0 0 0 0 0.4 0 0.84 0 0
Bolivina antiqua D’orBigny 2.23 0 0.41 0 0.75 0 0.41 0 0 0 0 1.14 6.85 7.02 0.49 0.45 0.75 0.41 0 0.4 0.48 0.42 0 0
Bolivinadilatatadilatatareuss 6.69 12.6 17.6 0 2.64 0.42 7.76 22.9 3.76 4.6 0.4 5.3 0 6.14 0.49 2.23 0 0.82 1.71 0.8 0 3.35 14.4 1.5
Bolivina hebes macfaDyen 1.12 0 0 2.29 0 0 0 0 0 0 0 0 0 0.44 0 0 0 0 0 0 0 0 0 0
Bolivinaplicatellacushman 1.86 4.6 1.23 0.76 0.38 0 0.41 0.8 0 0 0 0.38 12.5 3.51 0 2.23 1.13 0.41 0.43 1.2 0 0 3.6 1.87
BolivinapokornyicichaetzaPletalova 0.37 0 1.64 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
Bolivinascalprataretiformiscushman 0 0 0 0 0 0 0 0 0 0 0 0 0 6.58 0 0 0 0 1.28 0 0 0 0 0
Buchnerinabuchneri(margerel) 1.49 3.77 2.05 1.53 0.75 0 1.22 2.81 0.47 0 0.4 0.38 2.42 3.51 0 0.45 0 0.41 1.71 0.4 0 0 0.45 0.37
Bulimina elongataD’orBigny 3.35 3.77 2.05 0.38 1.51 1.26 5.31 5.22 4.23 4.18 1.21 4.17 0.4 0 0 0.89 2.63 0.41 0.85 3.61 0 1.67 4.05 2.62
Bulimina striata D’orBigny 2.23 2.09 0.82 0.38 0.75 0 1.22 1.2 0 0.84 0 1.14 2.02 2.19 0.49 0 4.89 1.65 0 2.41 0 2.93 0.9 2.62
Cancris auriculus (fichtel et moll) 0 0 0 0 0 0.42 0 0 0 0 0.4 0 0 0.44 0 0 0.38 0 0 0 0 0 0 0
CassidulinalaevigataD’orBigny 4.83 2.09 3.28 0 0.75 1.26 9.39 10.8 2.35 3.77 0 9.09 4.84 6.14 0.49 0.89 1.13 0.82 1.71 0.4 0 0.42 2.25 0
Cibicides sp. (small-sized) 4.83 9.62 16 8.78 6.79 2.52 12.7 14.5 16 5.02 10.1 18.9 16.9 13.6 3.9 13.4 0.75 2.06 11.5 7.63 1.9 18.8 0 0.37
Cibicidoides austriacus (D’orBigny) 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
Cibicidoides ungerianus (D’orBigny) 0.74 1.26 2.87 3.05 2.26 2.94 2.86 2.01 3.29 2.09 0 3.79 1.61 3.51 0.49 3.57 1.5 3.29 2.56 2.41 5.24 2.93 2.25 3.37
Dentalina sp. 0 0 0 0.38 0.75 0 0 0 0 0 0 0 0 0.44 0 0.45 0.38 0 0 0 0 0 0 2.25
Elphidiumcrispum(linne) 0 0 0 0 0 0 0 0 0 1.67 0.4 0 0 0 4.39 0 0.38 4.12 0.43 1.2 15.2 0 0 0
Elphidiumfichtellianum(D’orBigny) 0 0 0 0.76 0.38 0 0 0.4 0.47 0.84 2.02 0.76 0 0 1.46 0.89 0.75 0.82 0.43 0 0 0 0.45 0
Elphidium flexuosum (D’orBigny) 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
Elphidiummacellumfichtel etmoll 2.97 1.67 0 12.6 8.68 14.7 4.9 4.02 9.39 6.28 12.2 7.95 2.02 1.32 7.8 5.36 5.64 8.23 7.69 7.23 3.81 4.18 0.9 0
Elphidiumortenburgense(egger) 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
Elphidium rugosum (D’orBigny) 0 1.26 0 0.38 0.38 2.52 0.82 0.8 0.94 0.42 3.24 1.89 0 0.88 0.49 3.13 2.63 0.41 2.56 2.01 1.43 0.42 0 0
Elphidium sp. (juvenile) 0 0 1.64 0 0 0 0 0 0 0 0 0 0 0.44 0 0 0 0 0 0 0 0 0 0
Ehrenberginaserratareuss 0 0.42 0 0 0 0 0 0 0 0 0.4 0 0.4 0.88 0.98 0 0.38 0 0.43 0.4 0 0 2.7 3
Fursenkoina acuta (D’orBigny) 0.37 0 0.41 0 0 0 0 0.8 0 0.42 0 0.38 0 0 0 0 0 0 0 0.4 0 0 0 0
Chilostomellaovoideareuss 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.41 0 0 0 0 0 0.37
Globocassidulinaoblonga(reuss) 8.55 12.1 8.2 0 4.91 0 4.08 4.02 2.35 0.84 1.21 2.65 4.03 4.39 0 1.79 0.75 0.41 2.14 2.41 0 5.02 3.15 0.37
Globulina gibba D’orBigny 0.37 0 0 0 0 0.42 0.82 0 0 0.42 0 0.38 0 0 0 0 0.38 1.23 0.43 0.4 0 0 0 1.87
Globulina spinosa D’orBigny 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.41 0 0 0 0 0 0
Grigelis pyrula (D’orBigny) 0 0 0 0 0 0 0 0 0 0 0 0 0.4 0 0.98 0 0.38 0.41 0 0 0.95 0 2.25 1.5
Hansenisca soldanii (D’orBigny) 2.97 1.67 2.87 0.76 1.89 0.42 2.04 4.02 1.41 1.67 1.62 1.89 2.02 1.32 1.95 3.57 1.88 0.41 1.28 0.8 0 0.42 3.15 1.87
Hanzawaia boueana (D’orBigny) 1.12 1.26 0 0 1.51 0.84 0.82 0.4 0.47 0.42 0.81 0.38 0.81 1.32 0 1.34 1.5 0 0.85 1.61 0 2.09 4.5 4.12
Heterolepa dutemplei (D’orBigny) 4.09 2.93 2.05 1.15 5.66 1.26 7.76 3.21 11.3 15.1 5.67 2.65 1.21 0.44 8.78 1.34 8.27 3.7 4.27 5.22 1.9 2.09 1.8 3.37
Heterostegina sp. 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
Hoeglundina elegans (D’orBigny) 0.37 0 0.82 0.38 0 0 0 0 0 0 0 0 0 0 0 0.45 0 0 0 0 0 0.42 2.25 4.49
Karreriellachilostoma(reuss) 0 0 0 0 0 0 0 0 0 0 0 0 0 0.44 0 0 1.13 0.82 0.43 0 0 0.42 0.45 5.62
Laevidentalina elegans (D’orBigny) 0 0 1.64 0 0 0 0 1.61 0.47 0 0 1.52 4.03 0 0 0 0 0 0 0 0 0 0 3.75
Lagena striata (D’orBigny) 0 0 0 0 0 0 0 0.4 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
Lagena hexagona (Williamson) 0 0 0 0 0 0 0 0.4 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
Lenticulinacalcar(linne) 0 0 0 0 0 0 2.45 0 1.88 0 0 0 0.4 0.44 0.49 0 1.5 0 0 0 0 0 0 0.37
Lenticulina clypeiformis (D’orBigny) 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
Lenticulina inornata (D’orBigny) 0.37 0 0 0.38 0.75 0 0 0 0.47 0 0 0.76 0.4 0 0 0 0 2.06 0 2.41 0 1.26 0.45 6.74
Lenticulina orbicularis (d´orBigny) 0 0 0 0 0 0.42 0 0 0.47 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
Lenticulina sp. (broken or abraded) 0 0 0 0 0 0 0 0 0 7.95 1.62 0 0 0 0 0 0 0 0 0 0 0 1.35 0
Lenticulina sp. (juvenile) 0.74 0.84 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
Lenticulina vortex (fichtel et moll) 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
Lobatula lobatula (Walker et JacoB) 7.81 8.79 5.33 19.1 8.3 17.7 15.1 4.82 13.2 11.3 15.4 11 11.7 13.6 10.7 8.48 9.77 8.23 12 12.1 6.67 10.5 3.6 3
Marginulina hirsuta D’orBigny 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.75 0 0 0 0 0 0 0
The Badenian parastratotype at Židlochovice 195
Z2/16.9-17.0m
Z2/16.6-16.4m
Z2/15.9-16.0m
Z2/15.4-15.5m
Z2/15.1-15.2m
Z2/14.4-14.8m
Z2/14.1-14.2m
Z2/13.9-14.0m
Z2/13.8-13.9m
Z2/13.7-13.8m
Z2/13.3-13.4m
Z2/12.9-13.0m
Z2/12.2-12.3m
Z2/11.5-11.6m
Z2/11.0-11.2m
Z2/10.8-10.9m
Z2/9.9-10.0m
Z2/9.2-9.3m
Z2/8.8-8.9m
Z2/8.5-8.6m
Z2/8.3-8.4m
Z2/7.8-7.9m
Z2/6.9-7.0m
Z2/6.0m
Martinottiella karreri (cushman) 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
Melonis pompiloides (D’orBigny) 1.49 0.84 0.41 0 0.75 0.42 2.45 0.4 0.94 0.42 2.02 0.38 2.02 1.32 1.46 4.02 3.01 1.65 1.71 1.61 2.86 2.93 2.7 3.37
Nonion commune (D’orBigny) 1.86 3.35 2.46 0.76 1.13 0 2.04 1.2 4.23 3.77 1.62 2.27 2.02 0.88 0.49 1.79 0.75 2.06 0.85 1.2 0.48 2.09 7.66 10.5
Nonion sp. 0 0 2.46 0 0 0 0.82 2.41 0 3.77 0 0 0 0 0 0 0 0 0 0 0 0 0 0
Nodosaria hispida d´orBigny 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.4 0 0 3.6 1.12
Nummoloculina contraria (D’orBigny) 0 0 0 0 0 0 0 0 0 0 0 0 0 0 2.93 0 0 0 0 0 0 0 0 0
Oridorsalisumbonatus(reuss) 0 0.42 0 0 0 0 0 0 0.47 1.26 0 0 0 0 0 0 0.38 2.06 0 0 6.19 0.42 0 0
Neugeborina longiscata (D’orBigny) 0 0 0 0 0 0 0 0 0 0 0 0.38 0 0 0 0 0 0 0 0 0 0 0 0
Pappina parkeri (karrer) 0 0.42 0 0 0 0 0 0 0 0 0 0 0.81 0 0 0 0 0 0 0 0 0 0 0
Pararotalia aculeata (D’orBigny) 0 0 0 0 0 0 0 0 0.94 0.42 0.81 0 0 0 0.49 0 0 0 0 0 1.43 0 0 0
Planularia lanceolata (D’orBigny) 0 0 0 0 0 0 0 0 0 0 0 0 0.81 0.44 0 0 0 0 0 0 0 0 0 1.12
Plectofrondicularia digitalis (neugeBoren) 0.74 0.42 0.82 0 0.75 0 0 0.8 0 0.42 0 0.38 0.4 0 0 0.45 0.75 0 0 0.4 0.95 0.42 0.45 0
Praeglobobuliminapyrula(D’orBigny) 0 0 0 0 0 0 0 0 0 0.42 0 0 0 0 0 0 0.75 0 1.28 0 0 0 1.8 0.37
Porosononion granosum (D’orBigny) 4.46 3.35 3.69 0 0 1.68 0.82 0 0.47 0 0 0 0.81 0 0 0 0 0 0 0 0 0 1.8 0
Pseudotriloculina consobrina (D’orBigny) 0 0 0.41 0 1.89 0 0 0 0.47 0 0.4 0 1.21 0 0 0.45 0 0 0 0 0 0 0 0
Pullenia bulloides (D’orBigny) 0 0 0 0.38 0.38 0 0.82 0.4 1.41 1.67 0.81 1.89 0 0.88 0 0.89 1.5 1.23 1.28 2.01 0.48 0.84 2.7 5.24
Pyramidulina raphanistrum (linne) 0 0 0 0 0 0 0.41 0 0 0 0 0 0 0 0 0 0 0.41 0 0 0 0.42 0 0.37
Quinqueloculina buchiana D’orBigny 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.42 0 0
Quinqueloculina hauerina (D’orBigny) 0 0 0 0 0 0 0 0 0 0.42 0 0 0 0 0 0 0 0 0 0 0 0 0 0
Reussellaspinulosa(reuss) 1.12 0 1.64 3.44 1.89 1.26 0 0 0 0.42 0 0.76 0.4 1.75 0.49 1.79 1.5 0.41 1.28 1.61 0 2.51 0 0
Rosalina sp. (cf. semiporata (egger) 1.12 0 0 1.91 1.89 1.68 0 0 0 0 0 0 1.21 0.88 0 0.45 0 0 0.43 0 0 0 0 0
Semivulvulina deperdita (D’orBigny) 1.49 2.09 0 0 1.89 0.42 0 0.8 0.94 0.42 1.62 0 0 0 0 0 1.13 0 1.71 0.8 0 0.42 0 0
Sigmovirgulina tortuosa (BraDy) 0 0 0 0 0 0 0 0 0 0 0 0 1.21 0.44 0 0 0 0 0 0 0 0 0 0
Siphoninareticulata(czJzek) 0 0 0 0 0 0 0 0 0 0 0 0 1.21 0 0 0 0 0 0 0 0 0 0 0
Sphaeroidina bulloides D’orBigny 0 0.42 0.41 0 0.75 0 0 0 0 0 0 0 0.4 1.75 0.49 0.89 1.5 0 0.85 0.4 0 0 0.45 3.75
Spiroloculina sp. 0.37 0.84 0 0 0 0 0 0.8 0 0 0 0 3.23 0 0.49 0.45 0.75 0.41 0.43 0.4 0 0 0.45 0.75
Spirorutilus carinatus (D’orBigny) 0.74 0 0 0 0 0 0 0 0.47 0.42 0 0.38 0 0 0 0 1.13 0.41 0 0.4 0 0.84 0.45 0.37
Stilostomella adolphina (D’orBigny) 4.83 1.67 0 0 3.4 0 0 0 0 0.42 0.4 0 0 2.63 0.49 2.68 2.63 0.82 0.85 1.61 0.48 0.42 5.41 3.75
Stilostomella elegans (D’orBigny) 1.49 0.84 0 0.38 0 0 0 0 0.47 0.42 0 0 0.4 1.75 1.46 0 1.13 1.23 0 0 0.48 0.42 9.46 4.87
Textularia gramen D’orBigny 1.49 0 0 0.76 0.38 0 0.41 0 0.94 0.84 0.81 0.38 0 0 0 0 0 2.06 0 0 5.71 0.84 0 0
Textularialaevigata D’orBigny 0 0 0 0 0 0 0 0.4 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
Trifarina bradyi cushman 1.49 0 0 0 1.13 0 0.41 1.2 0 0 0 1.14 3.63 1.75 0 0.45 0 0 0 0.4 0 0 2.7 0.37
Uvigerina aculeata D’orBigny 0.74 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.42 0 0.75
Uvigerinabononiensisfornasini 1.12 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.75
Uvigerina macrocarinata PaPP etturnovsky 0 0 0 0.38 0.38 0 0 0 0 0 0 0 0.4 0 0.98 0 2.63 1.23 0 0.8 0 0 2.25 0
Uvigerina pygmoides PaPP etturnovsky 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
Uvigerina semiornata d´orBigny 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1.8 0
Valvulineria complanata (D’orBigny) 2.23 0 0 0 0 0 2.45 0 0 0 0 0 0 0.44 0 0 1.13 0 0 0.4 0 0 0 4.87
P/B 37.6 35.6 31.8 2.6 31.4 6.3 9.93 41 3.62 4.02 6.44 23.9 34 29.2 8.89 28.4 45.6 41.7 26.9 37.6 11 25.8 41 34.9
Foraminiferal number
(specimens/1 g of rock sample)
287 337 358 242 139 408 54.6 141 22.1 33.2 17.6 312 691 587 15 313 176 50.2 320 239 18.9 64.4 96.9 123
196 N. Doláková et al.
196N.Dolákováetal.
Appendix2:Listandrelativeabundancesofplanktonicforaminiferalspecies.
Z1/11.5-11.6m
Z1/10.2-10.3m
Z1/9.8-9.9m
Z1/9.0-9.1m
Z1/8.9-9.0m
Z1/8.5-8.6m
Z1/7.8-7.9m
Z1/7.2-7.3m
Z1/6.8-6.9m
Z1/6.3-6.4m
Z1/6.3m
Z1/6.0-6.1m
Z1/5.7-5.8m
Z1/5.1-5.2m
Z1/4.8-5.0m
Z1/4.3-4.4m
Z1/4.2-4.3m
Z1/4.0-4.2m
Z1/3.6-3.7m
Z1/3.1-3.2m
Z1/2.6-2.8m
Z1/2.2-2.3m
Z1/2.0-2.1m
Z1/1.9-2.0m
Z1/1.7-1.8m
Z1/1.2-1.3m
Z1/1.0-1.1m
Paleoecologicalcharacteristic(Reynolds &
Thunell 1985; hemleben et al. 1989; KelleR &
macleod 1992; Pujol & VeRgnaud gRazzini
1995;bicchietal.2003;sPezzafeRRi 1995;RuPP &
hoheneggeR 2008)
Turborotalitaquinqueloba(naTland) 0 21.7 34.2 0 55.2 6.06 14.6 18.2 28.8 7.69 7.08 0 5 0 0 0 38.9 0 34.8 34.2 45.2 2.08 5.66 0 20 31.3 47.4 Mixed wather column, (?)eutrophic, stress, tolerant
Globigerina ottnangiensis Rogl 9.52 3.11 0 0 0 0 4.85 5.61 6.21 15.4 2.65 0 25 0 0 0 0 25 0 3.36 8.1 4.17 0 11.6 8.82 5.36 2.26
Globigerina tarchanensis subboTina et chuTzieVa 0 11.2 18.7 4.48 19.5 0 3.88 17.8 0 0 0 6.67 0 0 0 0 41.9 0 28.3 29.5 27.6 8.33 11.3 3.16 17.7 35.7 25.6
Globigerina praebulloides blow 26.2 28.6 9.76 13.4 5.84 24.9 32 18.7 13 11.8 6.19 0 5 0 0 0 0 0 0 0 0 0 0 5.26 0 0 0
Globigerinabulloides d’oRbigny 14.3 1.24 4.88 8.96 0.65 21.8 12.6 7.48 14.7 4.73 7.08 0 0 0 0 16.7 6.59 25 17.4 6.04 6.67 27.1 35.9 34.7 16.5 10.7 3.01 Eutrophic, intermediate dweller
Globigerina bulloides d´oRbigny (with bulla) 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
Globigerinoides bisphericus Todd 9.52 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1.2 0 0 0 0 3.13 3.77 0 1.18 0 0 Oligotrophic, superficial dweller
Globigerinoides quadrilobatus d´oRbigny 2.38 0 0 2.99 0 0 0.97 0 1.69 1.78 0.88 0 0 0 0 0 0 0 0 0 0 1.04 0 0 0.59 0.89 0 Oligotrophic, superficial dweller
Globigerinoides trilobus (Reuss) 0 0 0 0 0 0 0 0 1.13 7.1 4.42 6.67 0 0 0 0 0 25 0 0.67 0 3.13 0 9.47 0 0 3.76 Oligotrophic, superficial dweller
Praeorbulinacircularisblow 7.14 0 0 2.99 0 1.82 0 0.93 0 0.59 1.77 0 0 0 0 16.7 0.6 0 0 0.67 0 3.13 0 0 0 0 1.5 Oligotrophic, superficial dweller
OrbulinasuturalisbRonnimann 0 0 0 0 0 0 0 0 0 0 0 13.3 0 0 0 16.7 0 0 0 0 0 5.21 0 0 0 0 0 Oligotrophic, superficial dweller
Globigerinella regularis (d’oRbigny) 7.14 3.73 8.13 13.4 2.6 3.03 4.85 1.87 6.78 12.4 11.5 40 20 0 0 0 1.8 25 2.17 0.67 0.95 19.8 7.55 5.26 1.76 2.68 2.26 Oligotrophic
Paragloborotaliaacrostoma(wezel) 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
Paragloborotalia mayeri (cushman et ellisoR) 23.8 30.4 24.4 53.7 16.2 42.4 26.2 29.4 27.7 38.5 55.8 26.7 40 0 0 0 7.78 0 17.4 20.8 10 15.6 32.1 22.1 32.4 13.4 10.5 Superficialdweller
Globorotaliaperipherodondablow et banneR 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Intermediate dweller
Globorotalia bykovae (aisensTaT) 0 0 0 0 0 0 0 0 0 0 2.65 6.67 5 0 0 50 1.2 0 0 4.03 1.43 7.29 3.77 8.42 1.18 0 3.76 Intermediate dweller
Catapsydrax sp. 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
Globoquadrina altispira (cushman et jaRVis) 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
Z2/16.9-17.0m
Z2/16.6-16.4m
Z2/15.9-16.0m
Z2/15.4-15.5m
Z2/15.1-15.2m
Z2/14.4-14.8m
Z2/14.1-14.2m
Z2/13.9-14.0m
Z2/13.8-13.9m
Z2/13.7-13.8m
Z2/13.3-13.4m
Z2/12.9-13.0m
Z2/12.2-12.3m
Z2/11.5-11.6m
Z2/11.0-11.2m
Z2/10.8-10.9m
Z2/9.9-10.0m
Z2/9.2-9.3m
Z2/8.8-8.9m
Z2/8.5-8.6m
Z2/8.3-8.4m
Z2/7.8-7.9m
Z2/6.9-7.0m
Z2/6.0m
Turborotalitaquinqueloba(naTland) 18.5 30.3 65.8 0 31.4 0 18.5 26 37.5 0 5.88 25.3 28.1 26.6 0 25 3.59 9.2 15.6 19.3 0 21.7 18.2 0
Globigerina ottnangiensis Rogl 3.09 6.06 0 0 0 0 0 0 0 0 0 0 0 1.06 0 0 4.93 5.17 2.46 3.33 11.5 0 0 2.1
Globigerina tarchanensis subboTina et chuTzieVa 11.7 15.9 27.2 0 15.7 0 0 38.7 0 0 41.2 26.5 44.5 43.6 0 8.06 0 0 10.7 0 0 0 0 0
Globigerina praebulloides blow 8.64 18.2 1.75 71.4 4.96 37.5 11.1 5.2 0 30 0 12.1 7.81 0 40 27.4 27.4 20.1 19.7 20.7 23.1 32.5 7.79 21.7
Globigerinabulloidesd’oRbigny 9.26 0 0 0 5.79 12.5 25.9 0 12.5 30 23.5 0 0 4.26 15 4.84 20.2 12.6 3.28 14.7 30.8 0 8.44 18.2
Globigerina bulloides d´oRbigny (with bulla) 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
Globigerinoides bisphericus Todd 0 0 0 0 0.83 0 0 0 0 0 0 0 0.78 0 0 0 0 0 0.82 0 0 0 0.65 1.4
Globigerinoides quadrilobatus d’oRbigny 1.23 0 0 0 0 0 0 0 12.5 0 0 0 0 0 0 0 0 2.3 0 0 0 0 1.3 8.39
Globigerinoides trilobus (Reuss) 0 0 0 0 1.65 0 0 0 0 0 5.88 0 0 1.06 0 0 2.69 4.02 0 0 11.5 0 1.3 14.7
Praeorbulinacircularisblow 0 0 0 0 0.83 0 0 0 0 0 0 0 0 0 0 0.81 0.45 1.72 0 0 0 0 0 0.7
OrbulinasuturalisbRonnimann 0 0 0 0 4.13 0 0 0 0 0 5.88 0 0 1.06 0 0 0.45 0 0 0 3.85 0 0 0.7
Globigerinella regularis (d’oRbigny) 3.09 0 0 0 3.31 6.25 14.8 3.47 0 20 5.88 12.1 0 1.06 20 1.61 6.73 4.6 2.46 2 15.4 0 0 1.4
Paragloborotaliaacrostoma(wezel) 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
Paragloborotalia mayeri (cushman et ellisoR) 43.2 29.6 5.26 28.6 31.4 37.5 25.9 22.5 37.5 10 5.88 22.9 15.6 21.3 25 31.5 33.6 39.7 45.1 39.3 3.85 45.8 59.7 23.8
Globorotaliaperipherodondablow et banneR 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
Globorotalia bykovae (aisensTaT) 1.23 0 0 0 0 0 3.7 4.05 0 10 5.88 1.2 3.13 0 0 0.81 0 0 0 0.67 0 0 1.95 4.9
Catapsydrax sp. 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
Globoquadrina altispira (cushman et jaRVis) 0 0 0 0 0 6.25 0 0 0 0 0 0 0 0 0 0 0 0.57 0 0 0 0 0.65 2.1
Appendix3:Listandrelativeabundancesofcalcareousnannoplanktonspecies.
Z1/0.7-0.8m
Z1/1.0-1.1m
Z1/1.2-1.3m
Z1/1.7-1.8m
Z1/1.9-2.0m
Z1/2.0-2.1m
Z1/2.6-2.8m
Z1/3.1-3.2m
Z1/3.6-3.7m
Z1/4.0-4.2m
Z1/4.3-4.4m
Z1/4.8-5.0m
Z1/5.1-5.2m
Z1/5.7-5.8m
Z1/6.0-6.1m
Z1/6.3-6.3m
Z1/6.3-6.4m
Z1/6.8-6.9m
Z1/7.2-7.3m
Z1/7.8-7.9m
Z1/8.5-8.6m
Z1/8.9-9.0m
Z1/9.0-9.1m
Z1/9.8-9.9m
Z1/10.2-10.3m
Z1/10.3m
Z1/10.9m
Z1/11.1m
Z1/11.5-11.6m
Z1/11.8m
Remarks (paleoecological characteristics
accordingbeaufoRT&aubRy 1992;
wells & oKada 1997; floRes et al. 1997;
bollmann etal.1998;Kameo 2002;wade
& bRown 2006)
Coccolithuspelagicus (wallich)schilleR 19.7 19.6 24.2 45.6 8.68 17.1 15.5 29.2 14.3 10.9 2.56 14.3 18.4 2.56 11.6 13.1 5.94 9.58 32.6 14.4 19.8 10.5 41.8 14 39.7 12.1 7.28 7.44 22.4 9.68 High-nutrient
Reticulofenestra minuta RoTh 73.9 76.5 69.7 46.5 86.8 81.1 70.8 62.8 73.7 85.3 97.4 85.7 80.2 95.7 80.4 81.5 86.8 79.2 56 72.1 67.6 77.3 39 78.6 11 69.5 44.1 31 61.8 53 Stress-tolerant, euryhaline
ReticulofenestrahaqiibacKman 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 5.94 1.25 0 0 0 0 0 0 22.4 0 36.4 48.4 0 30
Reticulofenestra pseudoumbilica gaRTneR 0 0.71 0.41 1.77 0.91 0 5.31 3.59 2.76 0 0 0 0 0.85 1.79 1.36 0 2.08 4.85 3.15 3.15 3.64 4.78 0.44 10.1 6.73 2.3 4.55 4.82 0
Cyclicargolithus abisectus (mülleR) wise 0.38 0 0.41 0 0.46 0 0.44 0 0 0 0 0 0 0 0 0 0 0 0 1.35 0 0 0.8 0 0.46 0 0 0 0 0 Reworked - Oligocene
Reticulofenestra bisecta (hay, mohleR & wade) RoTh 0 0 0.41 0.88 0 0 0 0 0 0 0 0 0 0 1.79 1.36 0 0 0.44 0 0 0 1.99 0 0.91 0.45 0 0 0 0.92 Reworked - Oligocene
Reticulofenestradaviesi(haq)haq 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.4 0 0 0 0 0 0 0 Reworked - Oligocene
Cyclicargolithus floridanus (RoTh & hay) buKRy 0.38 0 0.82 0 0 0 0.44 0 0.92 0.78 0 0 0.94 0 2.68 0 0 0 2.64 0 0.9 0 2.79 0 0 0.45 0 1.24 1.32 0
Helicosphaera carteri (wallich) KamPTneR 1.89 0 0.82 1.33 0 0 0.44 0.9 0.92 0 0 0 0 0 0.89 0 0.91 1.25 0 0 2.25 0.45 2.39 1.31 1.37 3.14 0.77 1.24 2.63 0.92
Helicosphaera walbersdorfensis mülleR 2.27 1.42 1.64 1.33 2.74 0.9 4.87 0.9 4.15 2.33 0 0 0.47 0 0.89 2.71 0.46 4.58 1.76 7.21 3.6 6.36 4.38 4.37 9.59 4.04 7.66 4.55 5.7 2.76
Syracosphaerapulchralohmann 1.14 1.42 0 0.44 0 0.9 0.88 0.9 1.84 0 0 0 0 0 0 0 0 1.25 0 0 0.45 0.45 0 0.44 0.91 1.79 1.53 1.24 0 0.46
SphenolithusheteromorphusdeflandRe 0 0 0.41 0 0.46 0 0 0.9 0 0 0 0 0 0 0 0 0 0.42 0 0 0.45 0 0.4 0 0.91 0 0 0 0 0.46
Sphenolithusmoriformis(bRonnimann &sTRadneR)
bRamleTTe & wilcoxon
0 0 0.82 0 0 0 0 0.45 0.46 0.78 0 0 0 0 0 0 0 0 0.44 0.9 0.45 0.91 0.4 0.44 0 0 0 0 0 0
Pontosphaeramultipora (KamPTneR)RoTh 0.38 0.36 0.41 1.77 0 0 1.33 0 0.92 0 0 0 0 0 0 0 0 0.42 0.88 0.9 0 0 0.8 0.44 0.91 0 0 0 1.32 0
Discoaster variabilis maRTini & bRamleTTe 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.44 0 0 0 0 0 0 0 0 0.41 0 0
Discolithinalatellipticabáldi-beKe&báldi 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.46 0.9 0 0 0 0.46 Reworked - Oligocene
Braarudosphaerabigelowii(gRan &bRaaRud)
deflandRe
0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.46 Euryhaline
Thoracosphaera spp. 0 0 0 0 0 0 0 0 0 0 0 0 0 0.85 0 0 0 0 0 0 0 0 0 0 0 0.9 0 0 0 0.46 Oligotrophic, stratified
Eifelithus spp. 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Reworked -Cretaceous, Eocene
Watzenauria spp. 0 0 0 0.44 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1.35 0.45 0 0 1.37 0 0 0 0 0.46 Reworked - Cretaceous
Nannoplankton abundance (semiquantitative, for
explanantion see Method)
vi vi v v v iv iv iii iv iii i i iv ii i vi vi vi ii iv iv iii iii v ii v v iv iv vi
Z2/16.9-17.0m
Z2/16.6-16.4m
Z2/15.9-16.0m
Z2/15.4-15.5m
Z2/15.1-15.2m
Z2/14.4-14.8m
Z2/14.1-14.2m
Z2/13.9-14.0m
Z2/13.8-13.9m
Z2/13.7-13.8m
Z2/13.3-13.4m
Z2/12.9-13.0m
Z2/12.2-12.3m
Z2/11.5-11.6m
Z2/11.0-11.2m
Z2/10.8-10.9m
Z2/9.9-10.0m
Z2/9.2-9.3m
Z2/8.8-8.9m
Z2/8.5-8.6m
Z2/8.3-8.4m
Z2/7.8-7.9m
Z2/6.9-7.0m
Z2/6.5m
Z2/6.0m
Remarks (paleoecological characteristics according
beaufoRT & aubRy 1992; wells & oKada 1997; floRes
et al. 1997; bollmann et al. 1998; Kameo 2002, wade &
bRown2006;stratigraphicalrangegRadsTeinetal.2012)
Coccolithuspelagicus (wallich)schilleR 4.85 15.1 7.41 3.3 12.6 15.16 4.62 9.83 23.1 7.96 6.43 1.77 3.57 0.93 12.7 14.43 10.3 12.06 11.79 12.96 1.36 4.65 12.15 5.09 21.21 High-nutrient
Reticulofenestra minuta RoTh 79.3 67.3 66.67 94.34 81.4 81.97 50.4 46.15 28.0 79.65 85.7 84.07 60.2 79.44 67.4 40.21 45.4 23.05 36.59 48.61 59.09 82.95 80.97 85.65 0 Stress-tolerant, euryhaline
ReticulofenestrahaqiibacKman 8.81 5.31 16.67 0 0 0 42.0 42.74 42.6 9.29 3.57 7.08 33.4 16.36 15.5 34.36 37.1 56.74 42.68 30.09 31.36 0 0 1.39 69.26
Reticulofenestra pseudoumbilica gaRTneR 2.64 4.49 3.7 0.94 0 0.82 0.42 0 0 1.77 0.71 2.21 0.45 0.47 3.3 4.47 4.13 6.38 1.22 3.7 5.91 6.2 2.02 2.78 0
Cyclicargolithus abisectus (mülleR) wise 0 0 0 0 0 0 0 0 1.22 0 0 0 0 0 0 0 0.83 0 0 0 0 0 0 0 0.43
Reticulofenestra bisecta (hay, mohleR & wade) Roth 0 0 0 0 0 0 0 0 1.22 0 0 0 0 0 0 0 0 0 0 0 0 0 0.4 0 0 Reworked - Oligocene
Reticulofenestradaviesi(haq)haq 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.78 0 0 0.43 Reworked - Oligocene
Cyclicargolithus floridanus (RoTh & hay) buKRy 1.32 0.41 0.37 0 0 0 0 0.43 1.22 0 0 0.88 0 0 0 0 0 0 1.22 0 0 0.78 0 0 0 Reworked - Oligocene
Helicosphaera carteri (wallich) KamPTneR 0 1.63 1.48 0 0.9 0 0 0 1.22 0 0 0.44 0 0 0 2.41 0 1.42 1.63 0 0 0.78 1.62 1.85 0
Helicosphaera walbersdorfensis mülleR 1.32 2.04 2.22 0.94 2.71 1.64 1.68 0 0 0.44 2.14 0.44 2.23 0.93 0.94 2.75 1.24 0 2.85 2.31 0.45 3.1 1.21 2.31 3.46
Syracosphaerapulchralohmann 0 2.45 1.11 0.47 2.26 0.41 0.84 0.43 0 0.44 0 0 0 0.93 0 0.69 0.83 0 0.81 0.46 0.45 0 0.81 0.46 3.03
SphenolithusheteromorphusdeflandRe 0.44 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.81 0 0.43
Sphenolithusmoriformis(bRonnimann &sTRadneR)
bRamleTTe & wilcoxon
0 0.41 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.78 0 0 0.43
Pontosphaeramultipora (KamPTneR)RoTh 0.44 0.82 0.37 0 0 0 0 0 1.22 0.44 0 0 0 0 0 0.34 0 0.35 0.41 0.93 0 0 0 0.46 0
Discoaster variabilis maRTini & bRamleTTe 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.43
Discolithinalatellipticabáldi-beKe&báldi 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.34 0 0 0 0 0 0 0 0 0 Reworked - Oligocene
Braarudosphaerabigelowii(gRan &bRaaRud)
deflandRe
0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Euryhaline
Thoracosphaera spp. 0 0 0 0 0 0 0 0 0 0 1.43 1.33 0 0.93 0 0 0 0 0 0.93 0.91 0 0 0 0 Oligotrophic, stratified
Eifelithus spp. 0 0 0 0 0 0 0 0 0 0 0 0.44 0 0 0 0 0 0 0.41 0 0 0 0 0 0 Reworked - Cretaceous, Eocene
Watzenauria spp. 0.88 0 0 0 0 0 0 0.43 0 0 0 1.33 0 0 0 0 0 0 0.41 0 0.45 0 0 0 0.43 Reworked - Cretaceous
Nannoplankton abundance (semiquantitative, for
explanantion see Method)
iii v v iv iv v iv iv i ii i iii vi v ii v vi v v iv iii ii v v v
TheBadenianparastratotypeatŽidlochovice197
198 N. Doláková et al.
Appendix 4: List and relative abundances of mollusca.
ŽIDL- 1 0.9-1 1.4-1.5 2.3-2.4 2.7-2.8 3.7-3.8 4.2-4.3 5.5-5.6 7.2-7.3 7.8-7.9 8.4-8.5 9.2-9.3
10.1-
10.2 11.9-12
BIVALVIA
Cubitostrea digitalina (DuBois, 1831) 1 2 1 1 1
Ostreidae indet. 2 1 1 1 2 1 2
?Pododesmus striatus (Brocchi, 1814). 1 1
Crassadoma multistriata (Poli, 1795) 2 2 2 1 1 2 2
?Aequipecten sp. 2 2 1 1 1
Costellamussiopecten cf. spinulosus (münster, 1833) 2 2 1 1 1
Aequipectencf.scabrellus(lamarck,1819) 1 1
Pectinidae indet. 2 1 1 2 2
Cardiidae indet. 1
Carditidae indet. 1
Veneridae indet. 1
Nuculanacf.fragilis(chemnitz,1784) 1 1 1 1
Nuculana sp. 1
Nucula sp. 1
Bivalvia indet. 2 2 2 1 1 2 2 1 1
GASTROPODA
Bittium sp. 2 1
Alvania sp. 1 1
Solariorbis sp. (cf. woodi (hörnes, 1856) 1
Gastropoda indet. 2
Cirripedia indet. 1 1
ŽIDL- 2 8.1-8.2 8.7-8.8 9.7-9.8
10.8-
10.9
12.7-
12.8
14.3-
14.4 15.9-16 16.9-17
BIVALVIA
Cubitostrea digitalina (DuBois, 1831) 1 1 2 1 1
Ostreidae indet. 2 2 2 2 2 1 1 1
Ostrea sp. 1
?Pododesmusstriatus(Brocchi,1814) 1 1 1
Crassadoma multistriata (Poli, 1795) 2 2 2 2 2 2 2 2
Aequipecten cf. macrotis (soWerBy in smith, 1847) 1 2 2 2 1 1
Aequipectencf.scabrellus(lamarck,1819) 1
?Aequipecten sp. 1 1
Costellamussiopecten cf. spinulosus (münster, 1833) 2 1 1
Flexopectencf.scissus(favre,1869) 1 1 1
?”Chlamys” trilirata (almera et Bofill, 1897) 1 1 1
?Macrochlamis sp. 1
Pectinidae indet 2 2 2 3 2 2 1 1
Cardiidae indet. 1
Veneridae indet. 1
Nuculana cf. fragilis (chemnitz, 1784) 1
?Spondylus sp. 1
?Corbula sp. 1
Bivalvia indet 2 3 2 1 2
GASTROPODA
Bittium sp. 3 1 1
Alvania sp. 3
?Gibbula sp. 1
Rissoina sp. 1
Solariorbis sp. cf. woodi (hörnes, 1856) 1
Gastropoda indet. 3 2
Cirripedia indet. 1 1
semiquatitative abundance (number of fragments): 1 - rare (1-5 fragments); 2 - common (6-15 fragments); 3 - abundant (over
15 fragments)
The Badenian parastratotype at Židlochovice 199
Appendix 5: List and relative abundances of ichthyofauna.
Deaniasp.(tooth)
„g.Etrumeus”weileri
Vinciguerriapoweriae
Valenciennellustripunctulatus
Benthosemafitchi
Diaphusacutirostrum
Diaphuscahuzaci
Diaphushaereticus
Diaphuskokeni
Diaphusregani
Diaphustaaningi
Diaphussp.juv.corroded
Lampichthysschwarzhansi
Notoscopelusmediterraneus
Grammonussp.
Physiculusaff.huloti
Gadiculusargenteus
Gadomustejkali
Coelorinchussp.
Trachyrincusscabrus
Merlucciusaff.merluccius
Sparidaeindet.(tooth)
Brachydeuterussperonatus
Acropomatidaeindet.
Trachurusaff.picturatus
Lesueurigobiusex.gr.vicinalis
Deltentosteustelleri
Gobiidaeindet.juv.corroded
ŽIDL-2:
6.6-6.7 m +
7.6-7.8 m +
10.6-10.8 m +
11.6-11.8 m + +
14.6-14.8 m +
15.6-15.8 m + + + + + +
16.6-16.8 m + + + + + ? +
ŽIDL-1:
2.8-3.0 m + +
6.6-6.8 m + + + + +
7.5-7.8 m + +
8.6-8.8 m + + +
9.6-9.8 m + + + + + +
10.6-10.8 m + + +
clay pit + ? + + + + + + + ? + + + + + + + + + + + + + +
Appendix 6: Composition of studied limestone samples with identified facies.
Thin-section/components Corallines Bryozoa Molluska Foraminifera Echinoids Serpulids Unsorted Micrite Sparite Litoclasts Pores Facies
42139-11151 ŽIDL-2 13.3 m 27.31 3.41 1.46 2.44 0.98 0 41.95 11.22 5.37 5.85 0 CAdl
42139-11251, ŽIDL-2 13.3 m 20.3 15.35 0 0.99 0 0 36.14 25.25 0.5 1.49 0 CAdl
42139-111051, ŽIDL-210.6 m 35.61 1.95 0.98 0 0 0 27.32 1.95 14.15 18.5 0 CAdl
42139-11351, ŽIDL-2 9.6 m 48.81 13.27 1.42 0 1.42 0.47 12.32 19.43 0 2.84 0 Bbr
42139-11551, ŽIDL-1 6.3 m 18.36 13.53 0 0.97 1.45 2.42 42.51 13.4 7.25 0 0.48 RFgm
42139-11751, ŽIDL-1 6.3 m 45.06 30.2 0 0 0 0 15.35 7.43 1.49 0 0.5 RFgm
42139-11851, ŽIDL-1 6.3 m 49.76 14.43 0 0 0.5 0.5 17.41 15.92 1 0.5 0 RFgm
42139-11951, ŽIDL-1 6.3 m 45.55 13.37 0 0 0.99 1.49 22.77 12.38 0 3.47 0 RFgm
42139-111151, ŽIDL-1 6.3 m 57.71 38.31 0 0 0 0 0.5 3.48 0 0 0 RFgm
42139-11651, ŽIDL-1 4.7-4.8 m 13.8 17.24 0 0.49 0 0.49 28.8 4.43 14.78 18.23 2.46 CAdl
42139-11451, ŽIDL-1 4.5-5 m 12.5 1.92 0 2.4 1.44 0 50.96 12.2 8.17 10.58 0 CAdl
200 N. Doláková et al.
Appendix 7: List and relative abundances of palynomorphs.
ŽIDL-2 - 246 m a s.l. ŽIDL 1 - 230m a s.l.
Taxon/m 15.6-8 16 16 16.6-8 16.8 7 7.5-8 8.5 9.7 10 10.6-8 10.911.1 11.8 11.9
marine Dinophyta several types Types with branched projections x x xx xx xx x xx xx x x x x x x
other Dinophyta Ovoidites… several types x x xx .
Cyanophyta gen. indet. Sigmopollis laevigatoides krutzsch &Pacltová 1990 x x x
Chlorophyta Tasmanaceae Pterospermellaeisenack1972,PleurozonariaWetzel1933 x x x x x x x x x
Botryococcus Botryococcusbraunii kützing 1969 . x x . x x . . x x
kubic caves x x x x x x x x
Sporophyta
- „ - gen. indet. Toroisporis sp. 1
?Polypodiaceae gen. indet.
Laevigatosporites haardti (Potonié & venitz 1934) thomson &
Pflug 1953 1 1 1 2 3 2 1 2 1 2 2
Davaliaceae Davalia Verrucatosporites alienus (Potonié 1931) thomson & Pflug 1953 1 1
Dennstaedtiaceae Paesia Verrucatosporites favus (Potonié 1931) thomson & Pflug 1953 1 1 1 1 1 1 x
Gleicheniaceae gen. indet. Neogenisporisneogenicuskrutzsch1962 1
Lycopodiaceae Lycopodium Retitriletes sp. 1 1
Lygodiaceae Lygodium Leiotriletes wolffi krutzsch krutzsch 1962 1 1 1 1 1
Corrugatosporites multivallatus (thomson &Pflug 1953) PlanDerová
1990/microvallatus(krutsch1967)nagy1985
1
- „ - Leiotriletesmaxoideskrutzsch1962 1 1 1
Lygodiaceae gen. indet.
Leiotriletessp.,TriplanosporitessinuosusPflug1952exthomson
&Pflug 1953
1 1 1 1 1
Osmundaceae Osmunda Baculatisporitesprimarius(Wolff 1934)thomson &Pflug 1953 1 1 1 x 1 1 1
Pteridaceae gen. indet.
Undulozonosporitessemiverrucatus(krutzsch1967)stuchlik
2001 1
Pteridaceae Pteris
Polypodiaceoisporites muricinguliformis nagy 1959/corrutoratus
nagy 1985 3 3 1 2 2 3 7 5 5 1 5 1
Selaginellaceae Selaginella Echinatisporis miocenicus krutzsch & sontag in krutzsch 1963 1
Gymnosperms
Pinaceae Pinus haploxylon and sylvestris types 63 600 228 182 168 348 278 120 540 650 198 260 46 200 123
Cathaya Cathayapolleniteskrutzschi(sivak 1976)PlanDerová 1990 13 100 28 14 14 27 44 28 63 70 23 31 9 35 27
Keteleeria Keteleeriapollenitesdubius (chlonova 1960) sloDkoWska 1994 1 1 1 2 1 1 1
Picea Piceapollis sp. 4 9 3 4 4 2 3 4 3 5 1 2 3 2
Abies Abiespollenites sp. 1 1 2 2 1 1
Cedrus Cedripitesmiocaenicuskrutzsch 1971 2 2 2 5 5 4 2 5 3 1 4 2 4
Tsuga Zonalapollenitesmaximus(raatz1937)krutzsch 1971 2 1 6 1 2 2 1 1 5 1
Sciadopitaceae Sciadopitys Sciadopityspollenitesserratus(Potonié &venitz 1934) raatz1937 2 7 7 2 2 3 4 5 4 1 5 2 3 2
Taxodiaceae Taxodium, Glyptostrobus
type
Inaperturopolleniteshiatus (Potonié)thomson &Pflug; I. conce-
pidites(WoDehouse)krutzsch
1 21 21 23 20 12 7 46 8 32 3 14 14 50
Sequoia Sequoiapollenites polyformosus thierg. 2 1 3
Ephedraceae Ephedra Ephedripites div. fsp. 1 1 1
Angiosperms
Aceraceae Acer sp. Aceripollenites striatus (Pflug 1959) thiele-Pfeifer 1980 1 1 1 1 2
Aquifoliaceae Ilex
Ilexpollenitesmargaritatus(Potonié1931)raatz1937exPotonié
1960 2 1 1 1 1 2
- „ Ilexpollenitespropinquus(Potonié
1931)thiergart 1937 ex
Potonié 1960 2 3 3 2 2 3 3 3
Araliaceae Aralia
Araliaceoipollenitesedmundi(Potonié 1931)Potonié 151ex
Potonié 1960 1 1 1 1 2
Hedera Araliaceoipollenitesreticuloidesthiele-Pfeifer1980 1 1
Asteraceae gen. indet. Tubulifloriditesmacroechinatus(trevisan1967)nagy1985 1 1 1 1 1
gen. indet. Cichorieaciditesgracilisnagy1969(nagy1985)
Artemisia Artemisiapollenites sellularis nagy 1969 2
Betulaceae Alnus Alnipollenites verus (Potonié 1931 ex Potonié 1960) 2 4 8 2 2 2 6 3 7 2 4 1 2 1 4
Betula Betulaepollenitesbetuloides(Pflug1953)nagy1969 3 4 4 2 3 2 1 4 4 2 4
Carpinus Carpiniditescarpinoides(Pflug1953)nagy1985 1 2 2
Ostrya Ostryapollenitesrhenanus(thomson 1950)nagy 1969 1 1
Buxaceae Buxus Buxapollisbuxoideskrutzsch1966 2 1 1 1 2 1 2 2 1
?Caryophyllaceae gen. indet. Caryophyllidites microreticulatus nagy 1969
Caryophyllaceae,
Alismataceae
Stellaria, Alisma Minutipollisgranulatuskrutzsch 1966
Caryophyllaceae gen. indet. 17 8 15 12 12 1 6 8 8 10 3 5 8 1 2
Cercidiphyllaceae Cercidiphyllum
Cercidiphyllites minimireticulatus (trevisan 1967) ziemBińskatWorzyDło
1994
1 1 6 1 1 2 1
Cornaceae Cornoideae, Mastix-
ioideae
Cornaceaepollis satzveyensis (Pflug 1953) ziemBińska-tWorzyDło
1994
1 1 1 1 2 2 2 1 2
Corylaceae Corylus Triporopollenites coryloides Pflug 1 1
Cyrillaceae, Clethraceae gen. indet.
Tricolporopollenitesmegaexactus (Potonié 1931) thomson &
Pflug 1953 2 6 3 2 2 4 4 13 5 12 3 4 4 8
- „ - Tricolporopollenites exactus (Potonié 1931) graBoWska 1994 3 2 2 4 3 1 1 2 2 1 4 4
Calluna Ericipitescallidus(Potonié1931)krutzsch1970 1 1 2 2
Eucommiaceae Eucommia
Eucommioipollisparmularius(Potonié 1934)ziemBińskatWorzyDło
1994
2 1 1 2 2 1 1 1
Fagaceae Fagoideae
Tricolporopollenitespseudocingulum(Potonié 1931)thomson &
Pflug 1953 2 1 2 2 3 1 4 2 1
Trigonobalanus Fususpollenites fusus (Potonié 1931) keDves 1978 2
The Badenian parastratotype at Židlochovice 201
ŽIDL-2 - 246 m a s.l. ŽIDL 1 - 230m a s.l.
Taxon/m 15.6-8 16 16 16.6-8 16.8 7 7.5-8 8.5 9.7 10 10.6-8 10.911.1 11.8 11.9
Castaneoideae, Trigo-
nobalanopsis
Tricolporopollenitescingulum(Potonié 1931)oviformisthomson
&Pflug 1953
6 4 3 4 4 5 8 16 9 21 3 6 6 10
?Castaneoideae X
Lythraceae (Decodon)
Tricolporopollenitescingulum(Potonié 1931)pusillus thomson
&Pflug 1953
4 3 2 4 3 1 3 4 2 3 3 1 2 4
gen. indet.
Quercoidites henrici (Potonié 1931) Potonié, thomson, thiergart.
1950
4 1 3 3 1 2 2 1 2 3
gen.indet.
Quercoidites microhenrici (Potonié 1931) Potonié, thomson,
thiergart. 1950
7 20 14 8 12 14 14 31 7 21 1 8 7 25
- „ - Quercoiditessp.,Q. granulatus(nagy 1969) sloDkoWska 1994, Q.
asper (Pflug &thomson 1953) sloDkoWska 1994
3 5 5 5 8 9 3 8 5 6 2 1 5
Quercusilextype 3 1 8 7 4 9
Fagus Faguspollenitesverusraatz1937 3 2 3 3 2 1 4 3 1 3
gen. indet. Tricolporopollenites liblarensis (thomson 1950) graBoWska 1994 18 13 26 31 28 10 16 20 15 5 15 1 17 7 15
gen. indet. Tricolporopollenitesquisqualis (Potonié 1934) krutzsch 1954 11 2 7 3 8 4 1 3 2 3
Hamamelidaceae Liquidambar Liquidambarpollenitesstigmosus(Potonié1934)krutzsch1954 1 1 1 1 1 1 2 3 3 1 3
Periporopollenites orientalis (nagy 1969) kohlan-aDmska &
ziemBińska-tWorzyDło 2009 3 2 1 1
Parrotia-Distylium
type
Tricolporopollenitesindeterminatus (romanovicz) ziemBińskatWorzyDło
1994
1
Parrotia-Distylium
type
Tricolporopollenitesstarosedloensiskrutsch &Pacltová1969
Chenopodiaceae gen. indet.
Chenopodipollis multiplex (WeylanD & Pflug 1957) krutzsch
1966 3 2 2 1 1 2 1
Juglandaceae Carya Caryapollenitessimplex(Potonié 1931)raatz1937 3 6 5 3 3 1 7 8 12 4 3 7 21 4 5
Pterocarya Pterocaryapollenitesstellatus(Potonié1931)thiergart 1937 1 1 2 1 2 3 2 1 1 2 1 1
Juglans Juglanspollenitesverusraatz1937 4 9 5 4 4 5 1 4 4 1 1 4 2
Engelhardia
Engelhardtioiditespunctatus (Potonié 1931) Potonié 1951 ex
Potonié 1960, E. quietus (Potonié 1931) Potonié 1951 12 21 25 15 12 20 14 29 8 33 1 25 16 18
Platycarya Platycaryapollenites miocaenicus nagy 1969 6 3 8 6 7 4 8 7 3 5 2 4 5 7
Lamiaceae gen. indet. 1 4 1
Lavandula t. 2
Salvia t. 2 2
Lythraceae gen. indet. Lythraceaepollenites sp. 1 4 1 3 2 1 6
?Magnoliaceae gen. indet. Magnoliapollisneogenicuskrutzsch1970 1 1
Myricaceae Myrica
Myricipitesbituitus(Potonié1931)nagy 1969/coryphaeus(Potonié
1931) Potonié 1960 7 15 14 9 11 13 17 21 8 17 4 6 26
Nyssaceae Nyssa Nyssapolleniteskruschi(Potonié 1931)nagy 1969 1 1 1
Nymphaeaceae Nymphaea Nymphaeaepollenites nagy 1969 1 1
Oleaceae Olea t. Oleaidearumpollenites sp. 15 9 14 18 15 2 8 20 11 13 4 8 7
Sambucus, Fraxinus Oleaidearumpollenitesmicroreticulatusthomson &Pflug
Fraxinus Fraxinoipollenites sp. 5 3 6 5 5 8 11 7 4 15 5 4
Plantaginaceae Plantago Plantaginacearumpollenitesmiocaenicusnagy 1 1 1
Platanaceae Platanus Platanipollis ipelensis (Pacltová) graBoWska 13 6 9 16 16 6 17 19 8 14 3 1 3 6
Polygonaceae Rumex Rumex t. 1 1 2 2 2 2 1
Ranunculaceae Thalictrum Thalictrum t. 1 1 1 4 2 3
Rosaceae ? Sorbus Sorbus t.
cf. Rubus type Rubus t. 1 1 1 1 1
Rubiaceae Galium Galium t 3 2 1 5 1 2 4 2
Rutaceae gen. indet Rutacearumpollenites sp. 1 2 3 1 2 2 3 1
Salicaceae Salix Salixipollenites sp. 7 1 1 1 3 4 3 1 2 1 3
Sapotaceae gen. indet. - several
types
Sapotaceoidaepollenites div. sp. 2 2 1 1 1
Staphyleaceae Staphylea Staphylea t. 1
Sterculiaceae Reevesia Reevesiapollistriangulus(mamczar1960)krutzsch 1970 1 1 1
Symplocaceae Symplocos Symplocoiditesvestibulum(Potonié 1931)Potonié 1960 1 1 1
?Tamaricaceae ?Tamarix Tamarixpollenites sp. 1 1 4 1
Tiliaceae Craigia Intratriporopollenitesinsculptusmai 1961 1 2 1 1
Tilia
Intratriporopollenites instructus (Potonié 1931) thomson & Pflug
1953 1 2 1 1 1 1 1 2 1
Tricolporopollenites
indet.
gen. indet. - several
types
Tricolporopollenites indet. 9 15 10 12 8 10 9 27 5 18 6 6
Ulmaceae Ulmus Ulmipollenites undulosus Wolff 1934 19 23 23 18 17 29 30 35 7 8 8 18 10 14
Zelkova Zelkovaepollenites potoniei nagy 1969 2 1 2 1 1 1 2 4 2 2
gen. indet. 1
Celtis Celtipollenites sp. 17 18 13 19 19 2 7 20 15 5 4 11 2 13
Urticaceae gen. indet. Urtica t. 3 1
Verbenaceae gen. indet. Tricolporopolleniteswackersdorfensisthiele-Pfeifer 1980 1 1
Clerodendrum type Clerodendrumpollenites minimireticulatus skaWinska 1994 1
Vitaceae Parthenocissus Tricolporopollenitesmarcodurensisthomson &Pflug 1953 1
Liliopsida
Arecaceae gen. indet. Arecipitessp. 1 1 1
Calamus DicolpopolliskockeliPflanzl1956 1 2 1 2 1 1 1 2 2
Butomaceae Butomus Butomuspolenites sp.
Poaceae gen. indet. Graminides sp. 2 6 4 1 1 1 1 1 3 7
Potamogetaceae Potamogeton Potamogeton sp. 2 1 1
Sparganiaceae Sparganium Sparganiaceaepollenites sp. 1 1
202 N. Doláková et al.
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165
Geology
Important changes in the paleogeography of the Western
Carpathians can be documented during the Late Miocene.
The basins represent grabens or half grabens; partly „pull
apart“ basins along strike slipe zones, but mostly flexural
type basins without apparent brittle tectonics, except for
normal faults at the basins´ margins (Kováč 2000).
The Vienna Basin is situated within the AlpineCarpathian
mountain chain, between the Eastern Alps and
the Western Carpathians. It represents a polyhistoric basin
with Neogene to Quaternary sedimentary fill, deposited in
various types of basins in relation to the paleotectonic
development of the orogen. During the Late Miocene, due
to paleogeographic changes, the connection with the Pannonian
basin became gradually closed. Lake Pannon
retreated southwards, and the northern coast of the back-arc
basin was slightly elevated due to progradation of deltaic
and alluvial facies, especially in the lowlands. Lake Pannon
was continuously filled by sediments transported by rivers
from uplifting mountain chains. The sedimentary environments
changed from deep to shallow lake and deltaic environment,
followed by development of alluvial plains. Due
to Paratethys isolation, the salinity decrease led to the
development of a totally fresh-water environment by the
end of this period. (Kováč et al. 1998).
Material and methods
All the studied samples were pelitic, partly calcareous
sediments. The lignite layers have not been used in these
studies.
The pollen data come from the well-preserved and welldetermined
plant macrofossil samples of the E. Knobloch
collection deposited in the Moravian Museum Brno, (Postorna,
Dubnany, Moravske Nova Ves surface localities
– 13 samples), claypit Gbely, deep boreholes (Suchohrad
32, Suchohrad 38, Jakubov 54 – 30 samples), and six shallow
boreholes (28 samples) made by the interdisciplinary
research programme (archeology, natural environment) of
the Masaryk University Brno at the Slavic (Great Moravian)
settlement Pohansko near Břeclav.
For maceration of the samples, HCl, HF and heavy liquid
ZnCl2 were used. Pure glycerine or glycerine gels were
used mostly as the observation media.
The coexistence approach is an efficient and reliable
method for quantitative terrestrial palaeoclimate reconstructions
in the Tertiary. It is based on the assumption that
Tertiary plant taxa have similar climatic requirements to
their nearest living relatives. The aim of the coexistence
approach is to find for a given fossil flora, the climatic interval
in which all the nearest living relatives of the fossil flora
can coexist (Mosbrugger and Utescher 1997).
SBORNÍK NÁRODNÍHO MUZEA V PRAZE AC TA M U S E I N AT I O N A L I S P R AG A E
Řada B – Přírodní vědy • sv. 64 • 2008 • čís. 2–4 • s. 165–173 Series B – Historia Naturalis • vol. 64 • 2008 • no. 2–4 • pp. 165–173
PANNONIAN VEGETATION FROM THE NORTHERN PART OF VIENNA BASIN
NELA DOLÁKOVÁ
Institute of Geological Sciences,Masaryk University,Kotlářská 2,611 37 Brno,Czech Republic,e-mail:nela@sci.muni.cz
MARIANNA KOVÁČOVÁ
Comenius University,Faculty of Sciences,Department of Geology and Palaeontology,Mlynská dolina,
SK – 842 15 Bratislava,Slovak Republic,e-mail:kovacova@fns.uniba.sk
Doláková, N., Kováčová, M. (2008): Pannonian vegetation from the northern part of Vienna Basin. – Acta Mus. Nat. Pragae, Ser. B, Hist.
Nat., 64(2–4): 165–173. Praha. ISSN 0036-5343.
Abstract. The studied pollen assemblages come from the Pannonian (Tortonian and Early Messinian) sediments in the Moravian and Slovak
part of the Vienna Basin. Vienna Basin represents a pull-apart basin situated between the Eastern Alps and Western Carpathian mountain
ranges. Due to the paleogeographic changes and climatic oscillations during the Late Miocene, the number of thermophilous taxa decreased,
and some of them disappeared completely. The variable heights and forms of the uplifted mountain chains created ideal conditions for
mixed mesophytic forests and extrazonal vegetation (Cedrus, Tsuga, Picea). The swamp, riparian, often hydrophilous (Azolla, Nymphaea,
Potamogeton) and halophyte (Chenopodiaceae) plants represent coastal swamps, local lagoons, and marshlands. Occasional occurrences of
dinoflagellates indicate slightly higher salinities, whereas green algae of the genus Pediastrum represent freshwater environments. The
amount of herbaceous plants (Artemisia, Asteraceae, Lamiaceae, Polygonum, Daucaceae, Caryophyllaceae, Plantago) increased.
n Late Miocene, Palynology, Vienna Basin, Paleoclimate
Received August 8, 2008
Issued December 2008
166
Vegetation and climate
Detailed palaeontological study of the Late Miocene
sediments of the Vienna Basin were realized by Bůžek
(1962) at the locality of Poštorná. At different localities, systematic
macrofloristic studies were carried out by Knobloch
(1962, 1963, 1968, 1969, 1972, 1981, 1985). In these
studies Knobloch categorised the Pannonian and Pontian
vegetation of this area into the four different mutually associated
floristic biotopes – 1.vegetation of the open water
level (Azolla, Nymphaea, Trapa, Potamogeton, Ceratophyllum)
2. coastal and coastal rim plants (Carex, Scirpus,
Phragmites, Sparganium) 3. plants associated with brown
coal swamp forest (Glyptostrobus, Nyssa, Alnus, Byttneriophyllum,
Myrica, Acer), and 4. vegetation growing on the
moist inner land stands (Carpinus, Betula, Fagus, Liquidambar,
Ulmus, Platanus)
At the Poštorná locality, Gabrielová (1966) interpreted a
marshy environment during the Pannonian based on the
pollen data from the coal seam. Kalvoda (1979), Lázničková
(2006) and Doláková et al. (2006) identified many
herbaceous taxa. Konzalová (2005) carried out a detailed
palynological study of the Pannonian palynospectra. She
documented an aquatic environment with a mix of freshwater
and brackish plankton, coastal herbaceous plants,
swamp and riparian forest, mixed mesophytic forest, and
extrazonal vegetation. In the studied samples she found an
absence of Myrica compensated for by water plants frequently
showing higher inundation.
Based on the previously published research and our new
pollen data, it was possible to characterize vegetation
assemblages and climate during the Pannonian.
Due to paleogeographical changes and climatic oscillations,
the number of thermophilous taxa decreased during
this time span, and some of them disappeared completely
from this area (e. g. Sapotaceae, Palmae). Mostly broadleaved
deciduous elements of warm – temperate mixed
mesophytic forests such as Quercus, Celtis, Carya, Tilia,
Zelkova, Ostrya, Liquidambar, Carpinus, Betula, Juglans
dominate generally (Pl.1, figs 2, 10-14). Thermophilous
elements and mixtures of Engelhardia, evergreen Fagaceae
morphospecies Quercoidites microhenrici, and less frequently
Quercoidites henrici, Trigonobalanopsis, Symplocos,
Cornaceaepollis satzveyensis, Tricolpopollenites liblarensis
up to a maximum of 25 % were present (Pl I., figs 5-9).
The various heights and forms of the uplifted mountain
chains created ideal conditions for a higher presence of extrazonal
vegetation (Cedrus, Tsuga, Picea, Cathaya) (Pl.1,
figs 15,16) in the investigated area. Nevertheless, according
to Ferguson et al. (1998) Cathaya which grows nowadays at
high altitudes, was adapted in the Miocene to the lower
ground conditions with enough air humidity. The sharply
demarcated facies changing both in time and space in their
individual pollen spectra were created by intrazonal types
of vegetation or by high amounts of herbaceous plants.
They included marshes and coastal swamps with Taxodiaceae,
Nyssa, and bush woods with Myrica. Ilex and Sciadopitys
replenished the plant spectra in the floodplain lowlands.
The prevailing vegetation types were often riparian
forests with Alnus (up to 20%, often 4-porate pollen grains),
Salix, Pterocarya, Liquidambar, Betula (up to 14%), Fraxinus,
Platanus (Pl.1, figs 1-3). Polypodiaceae, Osmunda,
and some thermophile ferns (Lygodiaceae) occured in the
moist and shady places. Shrubs and lianas such as Buxus,
Ephedra, Ericaceae, Vitaceae, Lonicera, Rosaceae type
Rubus (Pl. 3, figs 15,16) occurred on drier substrates in the
associated riparian forests. Accumulations of Chenopodiaceae
in the interfluvial areas probably indicate local saline
swampy environments during falls in sea level. The
increased amounts of herbs (up to 30% in pollen spectra)
indicate the existence of local open places such as wet
meadows (Thalictrum, Rumex, Valeriana, Dipsacaceae,
Lamiaceae, Galium) or areas which were never inundated
(Artemisia – up to 17%, Asteraceae, Campanula, Fabaceae,
Daucaceae, Caryophyllaceae, Plantago – Pl. 3). Poaceae
could even have been components of the associations forming
the undergrowth of the forest margin.
Aquatic plants created belts in shallow waters (Nelumbo,
Nymphaea, Myriophyllum, Sparganium, Potamogeton –
Pl. 2.), and along the water/land boundaries (Decodon,
Polygonum persicaria, Caltha, Valeriana – Pl. 3). Very
interesting findings are represented by microsporangia –
massulae (within small circled microspores) with very characteristic
glochidia of the freshwater fern Azolla bohemica
(Pl. 2) described by Pacltová (1958). Isolated glochidia,
found separately, were also frequent. According to Knobloch
(1981) this genus occurs in eutrophic waters today.
Due to the absence of glochidia, part of microsporangia is
indistinguishable from the genus Salvinia. Similar results
were published by Konzalová (2005) from the locality of
Poštorná near Pohansko. Knobloch (1981) identified seeds
and fruits of both mentioned taxa at Pannonian – Pontian
localities within this area (Azolla – Dubňany, Čáry, and
Salvinia – Ořechov-Mistřín).
Occasional occurences of dinoflagellates and green
algae Tasmanaceae indicate a slightly higher salinity, Botryococcus
can thrive in both brackish or freshwater environments,
whereas green algae Pediastrum, Mougeotia, aquatic
ferns Azolla, and aquatic and coastal plants (Nelumbo,
Nymphaea, Myriophyllum, Sparganium, Potamogeton etc.)
represent freshwater environments.
The noticeable Pediastrum cenobia belong predominantly
to the species P. simplex and P. boryanum which are
typical for open waters of eu- to mesotrophic conditions
(Komárek and Jankovská 2001, Miola et al. 2006, Zetter
1987). The non-pollen palynomorph Sigmopollis occurred
commonly. This morphotype is very similar to the Quaternary
type 128 after Van Geel et al. (1983). In our samples it
was often accompanied by type 74 as referred to by the
same authors. Both these types possibly pertain to algal
palynomorphs according to Van Geel et al. (1983), and
Miola et al. (2006), indicating open water environments in
eu-/mesotrophic conditions. Observations under the fluorescent
microscope also support their determination as algal
spores. Due to the chemical differences in sporopollenin of
lower and higher plants, and also to different rates of corrosion,
the algal bodies show very high fluorescence intensities,
whereas other palynomorphs show much lower, and
fungal remains are completely invisible (Van Gijzel 1971,
Yeloff and Hunt 2005, Doláková and Burešová 2007).
The fact that the sea level fell at the beginning of the
Late Miocene and led to large-scale erosion of older sedi-
167
ments in the area of the back-arc basin system is documented
in the Early Pannonian pollen spectra, where a lot of
redeposited sporomorphs of subtropical and tropical ferns
appeared (Slamková 2004). A higher percentage of nonarboreal
pollen (10-14%) indicates local marshes and partly
open woodland vegetation. The increase in halophyte taxa
documents the presence of coastal swamps, local lagoons
and marshlands during the lowstand of the brackish sea
(Kvaček et al. 2006).
During the Late Pannonian, the Western Carpathian paleogeography
started to change. Lake Pannon retreated
southwards and the northern coast of the back-arc basin was
slightly elevated due to progradation of deltaic and alluvial
facies, especially in the lowlands.
Swamp vegetation with straight growth in the swamp
substratum is mainly characterized by Taxodiaceae trees.
They are often present in association with Myricaceae, less
often with Nyssaceae. The riparian forest elements subdominantly
occurred with Alnus and Ulmus, mixed mesophytic
forests with Carya, Quercus, Craigia, Carpinus, Fagus and
herbs were represented by Chenopodiaceae, Asteraceae,
Ericaceae, Poaceae and Artemisia. Extrazonal vegetation of
the mountain areas with Picea, Tsuga, Abies, Cedrus is
common in the pollen spectra.
Paleoclimatic data quantified by the Coexistence
approach method (Mosbrugger and Utescher 1997) characterized
a climate in several categories. Using primary pollen
data from the Pannonian sediments of the Slovak part of the
Vienna Basin, the mean annual temperature (MAT) was
between 15.6–21.7°C, the coldest month temperature
(CMT) between 5.0 13.6°C, the warmest month temperature
(WMT) between 13.8–27.9 °C, mean annual precipitation
(MAP) between 373.0– 520.0 mm, the wettest month
precipitation (WtMP) between 73.0– 45.0 mm, the driest
month precipitation (DMP) between 5.0– 9.0 mm, and the
warmest month precipitation (WMP) between 27.0–227.0
mm (Kováč et al. 2006).
Discussion
A temperate climate with broad-leaved deciduous and
warm – temperate mixed mesophytic forests, was interpreted
for all the areas adjacent to the Vienna Basin. Increasing
amounts of herbaceous plant pollen were also observed. It
was presumed by Utescher et al. (2000), that the gradual
cooling started from the 14 Ma untill the Late Pliocene and
seasonality increased from the beginning of the Late
Miocene Planderová et al. (1993a,b) noticed a clear floristic
differentiation in representation of paleotropical and arctotertiary
elements between the southern and northern part
of Central and Eastern Europe during the Pannonian.
The Danube Basin situated at the Alpine-CarpathianPannonian
junction represents a region of the Central
Paratethys, strongly influenced by the orogen building
processes and climatic changes (Kováč 2000). In the reference
section of the Tajná 1 borehole, in the Lower Pannonian
sequence, dominant vegetation was formed by the
swamp representatives Taxodiaceae – Myricaceae with subdominant
presence of Nyssa, Alnus, Carya, Quercus deciduous,
Engelhardia, Chenopodiaceae and Poaceae. In the
Middle Pannonian sequence, changes in proportion of the
dominant elements are apparently related to the mild cooling
of climate and beech has been partly supplanted by fir
and deciduous oak. Taxa ratio in the predominant association
changed. The proportion of beech in Abies-Quercus
(deciduous) – Fagus association decreased. Oleaceae, Myrica,
Carya, Pterocarya, Alnus, Nyssa, Picea, Tsuga and
Cedrus occurred subdominantly (Kováč et al. 2006). Presence
of an increasing number of coniferous taxa Picea,
Tsuga and Abies also observed in earlier studies of the Pannonian
sequences (Nagy and Planderová 1985), can be
interpreted as a consequence of two factors: higher relief in
the hinterland of the basin or transition to seasonality. From
the Danube Basin, Planderová (1972, 1984, 1990) described
reduced marshes, isolated lakes with floras surrounded
by steppe meadows with scarce woody plants. In comparison
with Hungary, the climate was cooler and drier with a
dominance of Artemisia over other herbaceous pollen
(Nagy 1985, Nagy and Planderová 1985, Planderová 1990)
during the Late Miocene.
Erdei et al. (2007) confirmed the significant role of paleogeography
– subsidence of the Pannonian basin – in the
appearance of Pannonian floras and vegetation types with
extremely low diversities. The authors characterised most
of the Pannonian localities by monotonous azonal swamp
associations with Byttneriophyllum predominating which
indicates warmer climatic conditions.
In Poland, Wazynska (1998), Sadowska et al. (1993)
and others presumed a temperate and relatively arid climate,
thus not stimulating the development of swamp forest
with Nyssa and Taxodium. They were replaced by moist
riparian forests with Alnus, Celtis and Pterocarya. More
arid terrains were occupied by mixed forests with large
amounts of conifers, especially pines, and with only scarce
paleotropical relics. The amount of herbaceous plant pollen
increased during this time span (Poaceae, Artemisia, the
family Asteraceae, Daucaceae etc.).
Pannonian (Meotian) pollen data from the Ukraine indicate
the development of steppe or forest-steppe areas with
Poaceae and Artemisia (Syabraj 2000, Syabraj et al. 2007).
Kovar-Eder (1987) analyzed Pannonian vegetation and
climate in the Central Paratethys region. She has established
that the percentage of evergreen species increased
towards the southeastern part of the investigated area, and
arguments for either xeromorphic mediterranean-like vegetation
or for steppe-like conditions are invalid.
Very rich pollen assemblages were determined from the
Late Pannonian sediments of the Styrian Basin (Hoffmann
and Zetter 2005). Six associated paleo-plant habitats were
distinguished by the authors in the ancient wetland system,
namely, belts of aquatic plants, freshwater marsh habitat,
clastic swamp habitat, natural levee or crevasse-splay habitat,
organic swamp and wet-prairie habitat. They identified
40 herbaceous taxa, which documented not only closed forest,
but also the herbaceous vegetation of more xeric layers.
Conclusion
Due to paleogeographical changes and climatic oscillations,
thermophilous taxa numbers decreased during the
studied time span, and some of them disappeared completely
from the northern part of the Vienna Basin. Based on the
168
macropalaeobotanical and pollen data, a temperate climate
with broad-leaved deciduous and warm-temperate mixed
mesophytic forests was interpreted. The marked facies
mutually changing in time and space in their individual
pollen spectra were created by intrazonal types of vegetation
(marshes, riparian, coastal and aquatic) or by high
amounts of herbaceous plants (existence of local open
places such as drier substrata in the associated riparian
forests, and wet meadows). Variable height and form of the
uplifted mountain chains created ideal conditions for a
higher presence of extrazonal vegetation.
Based on pollen data from the Pannonian sediments of
the Slovak part of the Vienna Basin quantified climatic data
(mean annual temperature, mean annual precipitation...)
were calculated.
Comparison with the adjacent areas confirms the existence
of a climatically dependent gradient between the
southern and northern part of Central and Eastern Europe
during the Pannonian as documented by floristic differentiation
in representation of paleotropical and arctotertiary elements.
Increasing amounts of the herbaceous plants pollen
were also observed.
Acknowledgements
The study has been sponsored by the Research Project
MSM0021622427 (Czech Republic), projects of the Slovak
Research and Development Support Agency APVV-51-
011305, APVV-0280-07 and European Science Foundation
EUROCORES programme TopoEurope project ESF-EC-
009-07.
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170
PLATE 1
171
PLATE 2
172
PLATE 3
173
Explanation to the plates
PLATE 1
1. four-porate Alnus – Alnipollenites verus POTONIÉ; Poštorná
3.
2. Betulaepollenites betuloides (PFLUG) NAGY; Pohansko
V9, 6 m.
3. Salixipollenites sp. + Alnipollenites sp.; Pohansko V9,
6 m.
4. Nyssapollenites rodderensis (THIERGART) KEDVES;
Pohansko V9, 4.2 m.
5. Engelhardia sp. – Engelhardtioidites quietus
(POTONIÉ) POTONIÉ; Pohansko V9, 3 m.
6. Tricolporopollenites liblarensis (THOMSON) GRABOWSKA;
Pohansko V7, 4.2 m.
7. Reevesiapollis triangulus (MAMCZAR) KRUTZSCH;
Moravská Nová Ves 7.
8. Quercoidites henrici (POTONIÉ) POTONIÉ,THOMSON
etTHIERGART; Pohansko V9, 3 m.
9. Platanus sp. – Platanipollis ipelensis (PACLTOVÁ)
GRABOWSKA; Pohansko V9, 3 m.
10. Sciadopitys sp. – Sciadopityspollenites serratus
(POTONIÉ et VENITZ) RAATZ; Pohansko V9, 6 m.
11. Zelkova sp. – Zelkovaepollenites potoniei NAGY;
Pohansko V9, 6 m.
12. Quercus sp. Quercoidites sp.; Pohansko V9, 6 m.
13. Quercus robur type; Pohansko V9, 3 m.
14. Juglans sp. Juglanspollenites verus RAATZ; Pohansko
V9, 3 m.
15. Picea sp – Piceapollis sp; Poštorná 3
16. Tsuga sp. – Zonalapollenites maximus (RAATZ)
KRUTZSCH; Pohansko V9, 6 m.
PLATE 2
1. Microsporangium Azolla bohemica PACLTOVÁ; Pohansko
V9, 6 m.
2. Isolated glochidia of Azolla bohemica PACLTOVÁ;
Pohansko V9, 6 m.
3. Micosporangium cf. Salvinia x cf. Azolla; Pohansko
V9, 6 m.
4. Pediastrum boryanum (TURP) MENEGH; Pohansko V9,
4.2 m
5. Pediastrum simplex MEYEN; Pohansko V3, 8 m.
6. Pediastrum boryanum (TURP) MENEGH var. boryanum;
Pohansko V3, 8 m.
7. Nelumbo sp. – Nelumbopollenites europaeus (TARASEWICH)
SKAWIŃSKA; Pohansko V9, 4.2 m.
PLATE 3
1–5. Artemisia sp. several types cf. Artemisiapollenites sellularis
NAGY; Pohansko V9, 3 m.
6. Plantago sp. – Plantaginacearumpollenites miocaenicus
NAGY; Pohansko V9, 3 m.
7. Centaurea jacea type – Pohansko V9, 3 m.
8.,9. Daucaceae gen. indet. several types; Pohansko V9,
3 m.
10. Caryophyllaceae gen. indet.; Pohansko V9, 3 m.
11. Asteroideae – Senecio type.; Pohansko V9, 3 m.
12. Asteroideae – Cichoreacidites gracilis NAGY; Pohansko
V9, 3 m.
13. Asteroideae – Tubulifloridites macroechinatus (TREVISAN)
NAGY; Pohansko V9, 4.2 m.
14. cf. Echinops type; Pohansko V9, 3 m.
15. Fabaceae gen. Indet; Pohansko V9, 3 m.
16. Rosaceae gen. indet.; Pohansko V9, 3 m.
17. Rosaceae – Prunus type; Pohansko V9, 3 m.
18. Thalictrum sp.; Pohansko V9, 3 m.
19. Humulus/Cannabis type; Moravská Nová Ves
20. Ranunculus type; Pohansko V9, 6 m.
21. Polygonum persicaria – Persicarioipollis persicarioidites
KRUTSCH; Pohansko V9, 6 m.
Palynology and natural environment in the Pannonian to Holocene sediments of
the Early Medieval centre Pohansko near Breclav (Czech Republic)
N. Doláková*, A. Roszková, A. Prichystal
Institute of Geological Sciences, Masaryk University, Kotlárská 2, 611 37 Brno, Czech Republic
a r t i c l e i n f o
Article history:
Received 29 June 2009
Received in revised form
4 May 2010
Accepted 22 May 2010
Keywords:
Early Middle Ages
Great Moravia
Pohansko
Palynology
Non-pollen objects
Carpinus
a b s t r a c t
Pohansko (Czech Republic) is an important Early Medieval centre of the Great Moravian Empire (9th
century AD). The locality has a settlement with archaeological findings from Mesolithic to the modern
times.
The studied sediments belong to the stream and flood deposits of the Dyje river, aeolian sands and
buried soils. The lower parts of some boreholes penetrated into the Upper Miocene sediments. 172
samples were palynologically analysed. The original forest with relative small human impact was
observed on the base of Holocene layers. Even sporadic Cerealia pollen were found here. A deforestation
was visible in the cultural layer; it is linked with existence of the Great Moravian agglomeration and its
fortification. Later, a partial forest reconstruction probably took place in the surroundings. Very abundant
pollen of human-exploited plants (cereals, herbs) were discovered in the filling of archeological feature
O1. The possible practical potential of rampart also as the flooding protection was indicated by the
existence of marshy plants palynomorphs. The existence of oxbow was proved in the excavated probe S3.
The lowermost sample was dated by 14C as 7830 Æ 60 BP. The findings of Carpinus pollen in the layer
dated as Early Atlantic support the earlier spreading of this genus in the Czech Republic (i.e. South
Moravia).
Ó 2010 Elsevier Ltd. All rights reserved.
1. Introduction
The archaeological locality Pohansko near Breclav lies approximately
2 km from the town of Breclav (South Moravia, eastern part
of the Czech Republic) (Fig. 1). It was a significant Early Medieval
centre in the core area of the Great Moravian Empire, 9th century
AD, interpreted as a munitio, emporium and palatium of the
Moravian Early Medieval rulers (Machácek, 2005). The site lies at
an altitude of about 155e157 m a. s. l. and is situated about 12 km
from the confluence of the Dyje and Morava rivers. Today the
surrounding area is woody and boggy with meadows and floodplain
forests.
One of the key problems of this Early Medieval fortified site (as
with other Early Medieval sites in South Moravia) that must be
solved in co-operation with geologists and palaeobotanists is
determining a conclusion about its decline. It is believed that at the
end of the 9th century AD, such environmental events occurred
that caused extensive flooding and the covering of the Early
Medieval sites by flood sediments.
Palynological studies have been carried out within a framework
of broad interdisciplinary (archaeological, geological, environmental)
research between the Faculty of Science and the Faculty of
Arts, Masaryk University.
2. Description of the studied area
2.1. Archaeology
Fifty-year-long ongoing archaeological research studies have led
to the discovery of settlement traces from prehistoric times
(Mesolithic) to the Middle Ages. Findings of microlithic stone
artefacts are typical for the Mesolithic age (Kalousek, 1966). The
Neolithic settlement in the above-mentioned locality was represented
by Linear Pottery culture. According to Podborský (1993),
the beginning of Linear Pottery culture in the Moravia area is dated
between 7650 and 7450 BP. More frequent are Eneolithic artefacts:
a fragment of greenschist axe (Dostál, 1975) and finds connected
with the Funnel Beaker culture and Channelled Ware. Even some
artefacts from the Bronze Age have been found here (Dresler, 2008).
According to a non-verified information, one burial-ground from
the Hallstatt period was found before systematic excavation.
Plentiful decorations from the La Téne Age are known to be
* Corresponding author.
E-mail address: nela@sci.muni.cz (N. Doláková).
Contents lists available at ScienceDirect
Journal of Archaeological Science
journal homepage: http://www.elsevier.com/locate/jas
0305-4403/$ e see front matter Ó 2010 Elsevier Ltd. All rights reserved.
doi:10.1016/j.jas.2010.05.014
Journal of Archaeological Science 37 (2010) 2538e2550
Fig. 1. 1. Localisation of the studied area. 2. Part of the geological map. 3. Aerial photo (after www.mapy.cz) of fortification with mark of the boreholes and profiles. 4. Extreme
flooding (2006) through the opening in the fortification e according to Machácek et al., 2007. 5. Kopcany e only the one preserved church from the Great Moravia Empire (9th
century). 6. Geological profile e according to Machácek et al., 2007. 1 e original surface 2 e Great Moravian rampart 3 e buried semiterrestric gleysols 4e6 e sandy-clay flood loams
7 e wind-blown sands (sand dune) 8 e grey fluvial fine sands and/or sandy clays 9 e weakly clay sands and sandy gravels 10 e coarse or middle-grained Upper Pleistocene fluvial
sands and gravels 11 e coarse-grained fluvial gravels 12 e blue-grey sandy clays and clays e Neogene 13 e boreholes.
connected to this area. Relics of the Roman age have also been left
here (Dostál, 1975). The youngest pre-Slavonic artefacts are
examples of pottery from the Migration Period. The time of the
greatest expansion is embodied by Pohansko from the period
900e1000 AD within the Great Moravian Empire (Machácek,
2005). After its destruction (Fig. 2), or since 1500 AD, there have
no longer been any settlements here.
2.2. Geological background of the locality
From a geological point of view, the Early Medieval Centre of
Pohansko lies within an extended flood plain near the confluence of
the Morava and Dyje rivers (Fig. 1). The flat valley around the ring
wall reaches a width of about 4 km and is filled with Holocene flood
loams. Towards the confluence, the width reaches up to 10 km. The
Fig. 2. 1. The inner part of the fortification. 2. The outer part of the fortification e destruction. 3. Excavated test trench S3 e sediments of oxbow, with 14
C dating. 4. Archaeological
feature O1 e Flood loams and clays intercalated with buried terrestric gleysols.
N. Doláková et al. / Journal of Archaeological Science 37 (2010) 2538e25502540
marginal slopes of the valley protrude some 5 m above the flood
plain and are composed of Middle Pleistocene (Riss) fluvial sandy
gravels with Late Würmian dunes of wind-blown sands. In some
places these dunes also protrude from under Holocene flood loams
in the flood plain; one of them was used to build the Early Medieval
fortified site. The dune could have originated in the Dryas III (Late
Würm), i.e. 1210e1700 BP (Havlícek, 2004). The sand dunes were
also partly resedimented in the Holocene, as is evidenced by some
buried Mesolithic chipped artefacts. The fine to medium grained
blown sands have a yellowish to brownish colour and, especially at
the dune base, alternate with coarser grained layers of what were
probably of fluvial genesis. These Quaternary fluvialeeolian rocks
form the uppermost part of the Cenozoic filling of the Vienna basin,
a large geological unit of the Outer Carpathians. The bedrock of the
Quaternary deposits in the area of Pohansko was obtained through
boreholes at various depths from 4.4 to 8.1 m and is represented by
grey clays of the Pannonian Age.
Originally the sand dunes had a height of between 6 and 8 m,
but recently because of younger flood loam deposits, they are only
1e2 m above the flood plain. Some lower dunes were even buried
under the flood loams. The beginning of flood loam sedimentation
is estimated to be about 3000e4000 BP at the initial phase of the
Subboreal period (Brízová and Havlícek, 2002; Havlícek, 2004;
Havlícek and Smolíková, 1994; Opravil, 1983b).
From the 10th and especially 12th century AD (Opravil, 1983b),
the more intensive sedimentation of flood loams precipitated (or
substantially contributed to) early medieval settlement at Pohansko
and other similar sites in the flood plain near the Morava and
Dyje confluence (Machácek et al., 2007).
The deposition of flood loams was not continuous and a few
hiatuses occurred. Pelísek (1942) describes two soil horizons within
the flood loam sequence in the area of the Lower Morava depression.
Opravil (1999) estimates their origin to be the end of the
Subboreal (i.e. about 2900 BP) and the end of the older Subatlantic
(around 1000 AD).
Flood loams and clays intercalated with buried terrestric gleysols
in the construction of the Great Moravian fortification are
recorded in Pohansko (Fig. 2) even before this time.
As both semiterrestric soils underlie the whole fortification area,
we can safely say that they pre-date the Great Moravian times. The
deposition of the youngest flood sediments, which covered the ruins
of the fortification from the outer side, took place after a short break
in deposition marked by the formation of humic horizon “A” of the
upper buried soil in the flood plain. The surface of this soil was active
during both the construction and destruction of the Great Moravian
rampart. After this intermezzo, the deposition of flood loams has
continued, with some interruptions, tothe present. This is evidenced
by the layer of the youngest flood clays, which covered the ruins of
the rampart from the outer side (Machácek et al., 2007).
2.3. Paleovegetation reconstructions based on previous
paleobotanical studies
Several authors such as Brízová and Havlícek (2002), Jankovská
et al. (2003), Opravil (1962, 1978, 1983b), Rybnícek and Rybnícková
(2001) and Svobodová (1990) have attempted to create reconstructions
of the vegetation composition of the South Moravian
regions near the confluence of the Morava and Dyje rivers that were
acquired on the basis of macroflora and palynology.
Rybnícek and Rybnícková (2001) presume the development of
grassy subxerophyle oak woods in the bottom lands of the Early
Atlantic (Neolithic) and the development of mixed linden-oak
woods with elms in the medially wet areas. There was a striking
human influence registered there. Due to progressive deforestation,
intensive agriculture and herding in pastures, humans changed the
mesoclimate during the Bronze and Older Iron Ages. They caused
the secondary development of stands of xerothermic vegetation
and, similarly, the development of peat in the valleys. Several
authors (i.e. Firbas, 1949; Lozek, 2007; Opravil, 1999) suppose that
there were transient climatic (primarily humidity) oscillations
during the whole younger Atlantic to Subboreal. The intensification
of precipitation in the Lower Subatlantic caused an increase in
erosion and the first levelling of the lower niches of the bottom
land, according to Lozek (2007) and Opravil (1999). Since then the
modern-day cultural landscape has gradually developed.
Opravil (1978, 1983a, 1999) made reconstructions of plant
assemblages in the surroundings of the Early Medieval Centre
(Great Moravia) Mikulcice on the basis of fossil macroflora. In the
more variegated morphology here, he noticed greater diversity and
other quantitative proportions in the fossil plants compared to
today’s. The surface of the bottom land was formed by fluvial sandy
gravel overgrown with sporadic vegetation before the main deposition
of the flood sediments. Moor vegetation, alders and the edges
of shrubby willows developed in the depressions along the water
streams and erosive trenches. Forest formations, primarily hard
flood-forest, grew in the elevated benches.
Svobodová (1990) conducted the first palynological analyses of
sediments from the Pohansko and Mikulcice localities. She documented
increased human impact in the profile: deforestation,
synanthropic elements (such as Artemisia, Plantago lanceolata,
Convolvulus arvensis, Centaurea cyanus, Viciaceae) and cultural
plants (Cerealia). She proved that the environment of Mikulcice had
the character of a town with a great agricultural foundation,
whereas Pohansko was surrounded by mixed oak wood (unfortunately
without any chronological or archeological support).
Findings of anthropogenic elements were documented by
Brízová and Havlícek (2002) from the underlying sediments of the
cultural layer of the Mikulcice fortification.
3. Methods
Overall, 172 samples were palynologically analysed. They came
from 3 profiles in the archaeological section R18 across the rampart
and its ruins, the sedimentary filling of archaeological feature (a pit)
O1, excavated test trench S3, separate samples from the sedimentary
filling of the rampart structure and 13 boreholes V1eV13 (to
a depth of 8 m) (Fig. 1). The studied Holocene sediments belong to
the stream and flood loams of the Dyje river, aeolian sands and
buried soils. The lower parts of the boreholes penetrated into
Pleistocene fluvial sandy gravels and the underlying clay sediments
of the Upper Miocene (Pannonian) age (Machácek et al., 2007).
The sediments contained a varied amount of organic matter.
Primarily soils and flooding loams had typically low proportions of
palynomorphs. During soil-forming processes, the oxidation and
intensive activities of Bacteria and Fungi can cause partial or total
decomposition of the organic walls of pollen grains and spores.
More resistant palynomorphs (Alnus, Tilia, Pinaceae) could be
overestimated in the pollen diagrams (Havinga, 1967). The flooding
sediments are deposited very quickly in a large capacity (mostly in
early spring), which means that the palynomorphs are very
dispersed in the mass of inorganic material.
The sediments were macerated using the HCl, HF, KOH and
acetolysis (Erdtman, 1960).
The application of heavy liquid (ZnCl2) was used to increase the
number of palynomorphs.
For the identification of palynomorphs, the following publications
were consulted: Beug (2004), Erdtmann (1957), Erdtmann
et al. (1961), Komárek and Jankovská (2001), Moore et al. (1991),
Reille (1995), Rybnícková (1974), Van Geel et al. (1983), Zetter
(1987).
N. Doláková et al. / Journal of Archaeological Science 37 (2010) 2538e2550 2541
The pollen diagrams were created using the POLPAL programme
(authors Walanus and Nalepka, 1999). Five samples were dated by
14
C dating in the Poznan Radiocarbon Laboratory (T. Goslar).
Some profiles and boreholes contained time-comparable lithological
(geologicepedologically defined) horizons (Figs. 1 and 2)
related to the development of the Dyje river bottom (see above).
Due to these facts, the vegetation history could be interpreted from
the most plentiful pollen spectra of different spots in the equivalent
layers. On this basis, two combined pollen diagrams were constructed
e one from the inner parts of the fortification (Figs. 1 and
5), another from outside of the rampart (Figs. 2 and 6).
We usually did not make pollen diagrams for each individual
plant, but instead plotted several plants into “ecological floral
units” to give a basic idea of the vegetation relationships. We realize
that there is some simplification: e.g Quercus (Opravil, 1978, 1983b)
could be a member of either mesophyte oak-lime-hornbeam or
flood-plain forest vegetation, depending on the species.
1. The unit “oak-lime-hornbeam forest” contains Quercus þ
Loranthus, Carpinus, Tilia, Acer (Fig. 3);
2. The group “Flood-plain forest” contains Alnus and Ulmus,
Fraxinus, Salix and Populus;
3. “Marshy and aquatic herbs” contains Potamogeton, Sparganium,
Filipendula, Myriophyllum, Nymphaea, Nuphar, Typha, Cyperaceae,
Caltha (Fig. 3);
4. Some primary as well as secondary anthropogenic indicators
are marked in yellow (Fig. 4);
5. All other herbs were included in the unit called “Other herbs”.
4. Results
4.1. The oldest sediments
The bottom sediments (6e8 m) were sedimentologically dated
as Upper Neogene-Pannonian (Machácek et al., 2007) (Fig. 1). The
pollen spectra with only Miocene palynomorphs (i.e. Engelhardia,
Symplocos, Glyptostrobus, Tsuga, Cedrus) (V9 e 6 m, V2 and V3 e
8 m) testified to these ages (Doláková and Kovácová, 2008).
Redeposited Miocene palynomorphs were also occasionally found
in some Holocene sediments (from a base of up to 1.5 m deep). The
preservation states of individual palynomorphs known both from
the Quaternary and Tertiary (e.g. Pinus, Ulmus, Alnus, Quercus,
Juglans) may be different or not under a light microscope. Observation
under a fluorescent microscope can help to determine the
reworked palynomorphs (Van Gijzel, 1971; Yeloff and Hunt, 2005;
Doláková and Buresová, 2007).
Particular coarse-grained Upper Pleistocene fluvial sands and
gravels, in positions similar to the boreholes (about 3,8e8.5 m)
(Fig. 1), represent the basal Quaternary layers (Machácek et al.,
2007) here and were palynologically sterile (due to their mechanical
properties).
4.2. The basal Holocene fluvial sands
Pollen spectra from fluvial fine sands and/or sandy clays correspond
to the former research of flood-plain vegetation near the
confluence of the Morava and Dyje rivers (see above).
The pre-domination of oak-lime-hornbeam forest (Quercus was
most frequent) alternating with flood-plain forest (dominant Alnus,
and less Ulmus, Fraxinus, Salix and Populus) was visible in the pollen
diagrams (Figs. 5 and 6), V5, V13 e outside of the ramparts (Fig. 6),
V1 (Fig. 7), P1, S3 (Figs. 2 and 8). This phenomenon could have been
caused by primeval human activity or climatic reasons. There was
regular occurrence of the pollen of Betula. Pinus sylvestris is
frequent; Picea was found in low quantities. The infrequent pollen
grains of Abies were transported here from the more distant higher
areas. The occurrence of Juglans pollen is very interesting and
disputatious. This genus had no natural occurrence in Central
Europe in the Holocene. Its cultivation by humans or re-deposition
from the underlying Miocene sediments should not be ruled out
(see discussion).
In addition, there is a minor proportion of shrubs (sporadically
Rosaceae e Prunus and Rubus); only Corylus was commonly found.
Trees dominated over herbs, but the landscape was not
completely forested. Associations of open locally moist areas were
represented mainly by the following herbs: grasses e Poaceae,
Asteraceae, Cyperaceae, Ranunculaceae, fewer Euphrasia, Chrysosplenium
and Symphytum. There were also plants growing at the
edges of bodies of water: Typha, Potamogeton, Nymphaea. Nevertheless,
there were some anthropogenic indicators in these layers e
archaeophytes such as Cerealia (Figs. 3 and 4) and field weed such
as Polygonum aviculare, C. cyanus (profile P1 e 2 m) (Fig. 3). There
were also secondary anthropogenic elements, original to this
landscape, but spreading thanks to human activity, which created
suitable places for them due to deforestation, ruderalisation,
pasturage and agriculture. They are as follows: P. lanceolata, Rumex
acetosella, R. acetosa, Artemisia (Fig. 3), and Chenopodiaceae.
These facts indicated that there was agriculture in the period
preceding Slavonic settlement, even in the time before the sedimentation
of the flooding loam in the bottom land.
A charcoal particle from a depth of about 5 m in borehole V13
was dated by 14
C as the 7350 Æ 50 cal BP (6370e6070 cal BC). The
14
C dating of overlaying sediment: 3 m: 8240 cal BP (7470e7070 cal
BC) proved the big sedimentological dynamics of the bottom land.
The Cerealia-type pollen (Cerealia X Glyceria species) from these
layers should be considered (see discussion).
4.3. The lower flooding sediments
A marked decrease in pollen grains of woody genera already in
the overlying lower flooding sediments and the lower buried soil
inside of them was documented. Their palynomorph content is
generally very low (see chapter 3.). Only a few samples yielded
more palynomorphs capable of creating an image of the vegetation.
Oaks (Quercus), hornbeams (Carpinus) and elms (Ulmus) almost
disappeared. Alders (Alnus) decreased and only poplars (Populus)
and pines (Pinus) remained. In contrast to the decrease of trees,
herbs (primarily grasses and ferns) increased (Fig. 5). It is presumed
that the great amount of deforestation allowed more flooding.
The regular occurrence of spores of Sphagnum, the increased
percentage of sedges (Cyperaceae) and the finding of aquatic plants
Myriophyllum spicatum (Fig. 7), Utricularia and green algae of
Botryococcus in the layers directly beneath and outside of the
fortification all provide evidence for the existence of marshes here.
The fortification must have been partially built on a swamp. It may
have also played a role in protecting against flooding. Practical
evidence of this was shown in 2006, a year of extreme flooding
(Fig. 1). The opening in the fortification for archaeological investigation
caused the inundation of a large part of the locality.
4.4. Sand dune
Borehole V1 (Fig. 7) was located on the sandy dune and contained
slightly loamy aeolian sands. Within the whole borehole, the
predominance of herbs over trees was noticeable (mainly Poaceae
and Asteraceae). P. sylvestris is the richest arboreal element. It is the
typical tree of sand dunes and also a significant pioneer tree,
penetrating open spaces such as fallow land first. It is often overvalued
in pollen spectra due to its strong resistance during
N. Doláková et al. / Journal of Archaeological Science 37 (2010) 2538e25502542
deposition (Havinga, 1967) or high pollen production and large
dispersion range. Tilia was more frequent here than in other
boreholes. This could be related to its better resistance in sandy
sediments. The number of elements of flood-plain forest was lower.
The alternating predominance of flood-plain forest over oak-limehornbeam
forest e similar to other boreholes and profiles e is
conspicuous. The pollen spectra indicated vegetation of drier places
with bad ground.
Fig. 3. Main pollen types, LM magnification 1000Â. 1. Quercus sp. e O1, 150 cm. 2. Tilia sp. e P1, 160 cm. 3. Acer sp. e V5, 320 cm. 4. Juglans sp. e V5, 320 cm. 5. Carpinus sp. e V5,
320 cm. 6. Carpinus sp.e S3, 135 cm. 7. Loranthus sp.e O1, 150 cm. 8. Galium sp. e O1, 154 cm. 9. Cerealia e Triticum type e V5, 320 cm. 10. Cerealia e Secale type e O1, 154 cm. 11.
Cyperaceae e P1, 200 cm. 12. Plantago lanceolata e V5, 250 cm. 13. Centaurea cyanus e P1, 200 cm. 14. Polygonum aviculare e V1, 230 cm. 15. Rumex sp. e V5, 250 cm. 16. Salix sp. e
P1, 200 cm. 17. Myriophyllum sp. e O1, 150 cm. 18. Sphagnum sp. e P3, 15 cm. 19. Botryococcus sp. e S3, 110 cm. 20. Probably aquatic moss, type 340 e S3, 110 cm. 21. Salvinia sp. e S3,
90 cm.
N. Doláková et al. / Journal of Archaeological Science 37 (2010) 2538e2550 2543
4.5. Excavated test trench S3
Most of the sediments were dark humic flooding loams (Fig. 2).
Palynology confirmed the sedimentological results regarding the
existence of an oxbow inside of the locality. Freshwater green algae
Pediastrum boryanum, frequent Botryococcus (Fig. 8) as well as
zygospores of Zygnemataceous algae, including Spirogyra and
Mougeotia, occurred in these sediments. There were plenty of nonpollen
types 128A and 74 (Van Geel et al., 1983), as well as some
possible algal palynomorphs. Sphagnum was sporadic, however,
there was an abundance of spores of type 340 found belonging to
the aquatic mosses, according to Miola et al. (2007) (Fig. 3).
Fig. 4. Author of SEM photos e J. Stelcl. 1e3. Daucaceae e Peucedanum sp. e O1, 150 cm 1. LM 1000Â magn. 2e3 SEM. 4e6. Quercus sp. e O1, 150 cm 4. LM magn. 1000Â, 5e6 SEM
7e9. Artemisia sp. e O1, 154 cm. 7. LM 1000Â magn. 8e9 SEM. 10e12. Poaceae e wild grasses e O1, 150 cm. 10. magn. 1000Â, 11e12 SEM. 13e15. Cerealia Triticum type e O1, 150 cm.
13. LM magn. 1000Â, 14e15 SEM.
N. Doláková et al. / Journal of Archaeological Science 37 (2010) 2538e25502544
Fig. 6. Combinated pollen diagram e boreholes outside of fortification.
Fig. 5. Combinated pollen diagram e boreholes and profiles from inner part of fortification.
N. Doláková et al. / Journal of Archaeological Science 37 (2010) 2538e2550 2545
Common sporangia with microspores of the water fern Salvinia
natans (Fig. 3) were found at some depths. There was also a regular
occurrence of water rim genera of Potamogeton, Typha, Sparganium
and the family Cyperaceae, sporadic M. spicatum and several
objects resembling Charophytes gyrogonia. These fossil types
indicate an open water environment in eu- to mesotrophic conditions.
From the uppermost part of the sediments, the water indicators
decreased.
The 14
C dating manifested surprisingly high ages of the dated
sediments (Fig. 2):
S3 1.27 m: 7830 Æ 60 BP (7050e6450 cal BC) as Lower
AtlanticeNeolithic;
S3 0.4 m: 2210 Æ 30 BP (380e190 cal BC) as Lower SubatlanticHallstatt;
La Téne.
The pollen diagram can be divided into two parts (Fig. 8). The
boundary is created by the practically sterile samples from the
middle part of the profile.
A pre-domination of trees over herbs was observed in the lower
part. In addition to Pinus, elements of oak-lime-hornbeam forest
also occurred. The pollen of Carpinus occurred sporadically, but it
was found at almost all depths from the lower part of the test
trench (even below the sample dated as Early Atlantic). These
findings are significant as they confirm the earlier spreading of
Carpinus (Figs. 3 and 6) in the Czech Republic and refute the
traditional assumption of their Subboreal penetration (see discussion).
Corylus reached 8% in some samples. Elements of flood-plain
forest prevailed in the lowermost part, where anthropogenic indicators
Cerealia and Juglans were found, although to a lesser extent
than in the upper part.
There were a visibly decreasing number of trees, increasing
number of herbs and more frequent primary and secondary indicators
of human activity (Cerealia, C. cyanus, P. lanceolata, P. aviculare,
Rumex acetosa, Chenopodiaceae) in the upper part of the studied
layers. The prevailing presence of flood-plain forest is visible at the
samples dated as Hallstatt, which correspond to the abundant
occurrence of Salvinia (see above in this chapter). A markedly
Fig. 8. Pollen diagram e excavated test trench S3.
Fig. 7. Pollen diagram of borehole V1 e through the sandy dune.
N. Doláková et al. / Journal of Archaeological Science 37 (2010) 2538e25502546
increasing amount of Pinus is noticeable in the uppermost sample,
compared to other boreholes and profiles (see below).
4.6. The cultural layer
A cultural layer of about 0.30 m was uncovered at a depth of
about 0.5 m from the surface in section R18 (Fig. 2). It represents
the homogenized upper buried soil with artefacts from the Great
Moravia age. Generally, herbs prevailed over trees (Fig. 9). The
base of the cultural layer probably occludes the natural floodplain
surface from the beginning of Early Medieval times. The
initial deforestation (mainly oak-lime-hornbeam forest group)
was connected with the existence of the Great Moravian
agglomeration and its fortification. High quality timber was
already consumed from the immediate vicinity of Pohansko (as
a building material for houses and fortification and as fuel).
Opravil (2000) analysed the coalified tree remains from the
construction of a protective wall. He concluded the following
composition: 75% Quercus, 5% Ulmus and 3% Fraxinus. Corylus also
had an increasing tendency. Juglans could have been cultivated by
the inhabitants of Pohansko. Herbs were evidently also abundant
(e.g. Asteraceae, Poaceae, Cyperaceae, Ranunculaceae, Rosaceae,
Rubiaceae).
Cereals were found in the whole cultural layer with the
maximum amount of 5%. Pollen grains of other cultural and synantropic
plants (e.g. Humulus-Cannabis, P. lanceolata, P. major/
media, Urtica) were found. Chenopodiaceae increased mostly in the
upper part of the cultural layer, which was related to ruderalisation
caused by human activity. The surroundings of the fortification
could have been used as pastures or fields.
4.7. Archaeological feature O1
Samples from the wall of archaeological feature (a pit) O1 were
almost sterile. Another two samples came from the anthropogenic
filling of this object. The lower samples were dated by 14
C as
2560 Æ 50 years BP (820e520 cal BC) as Lower Subatlantic e
Hallstatt.
The samples contained about 24% trees and 76% herbs (Fig. 9).
Elements of flood-plain forest slightly prevail over the oak-limehornbeam
forest. Pinus was the most abundant tree in both
samples. Abies was also more highly represented. We can assume
that it was carried by the wind from more distant regions. Abies was
used as a revetment of a grave in the nearby Great Moravia locality
Mikulcice (Opravil, 1962). It could have been that this precious tree
was imported on special occasions from more distant places.
In both samples Sambucus, which prefers moist nitrogenous soil,
was found. Juglans and Corylus may have been cultivated for nuts.
Herbaceous pollen was very abundant (Fig. 9). Meadow herbs
include Centaurea sp., Centaurea jacea, Euphorbiaceae, Euphrasia,
Fabaceae (Lotus, Trifolium), Rosaceae and Veronica. Plants such as
Daucaceae (Fig. 4), Fabaceae, Galium, Polygonaceae and Ranunculaceae
could belong to a category of herbs which grew near the
edges of woods or in damp places. There was a frequent occurrence
Fig. 9. Pollen diagram from the Great Moravian cultural layer and filling of the archaeological feature O1.
N. Doláková et al. / Journal of Archaeological Science 37 (2010) 2538e2550 2547
of marshy and aquatic herbs (Cyperaceae, Chrysosplenium, Myriophyllum,
Potamogeton/Sparganium).
In both samples, a prominently high amount (over 13%) of Cereal
(Triticum and Secale) types (Figs. 3 and 4) were found (Fig. 9). This is
evidence of the probable manipulation or processing of cereals in
human settlement. The following plants were connected to the
presence of human and agricultural activities: weeds such as
Polygonum persicaria and Silenaceae and other synantropic herbs
such as Urtica, Humulus/Cannabis, P. lanceolata, Plantago major/
media and P. aviculare.
We suppose that some plants could have been collected or
cultivated and used as medicinal plants (e.g. Sambucus, Euphrasia,
Salvia, Plantago, Alchemilla, Urtica, Valeriana, Artemisia). Some
plants could have been used for dyeing textiles (e.g. the fruits of
Sambucus and some Rosaceae, leaves of Urtica, Juglans and
Betula). It is necessary to support this hypothesis with further
research and evidence.
Both samples are different from the other samples from
Pohansko. The higher accumulation of normally scarce pollen could
be connected to human activity within the location of the fort.
4.8. Samples of sediments from the stone rampart
The pollen spectra were very poor. They contained almost all the
trees known from the other samples in this locality. Asteraceae
were the abundant herbs. There were many charcoal particles and
non-pollen objects. Most of them were spores of terrestrial algae
and spores and conidia of Fungi often living on decomposed wood
(some of them on oak). Sporadic cysts of marine algae originated
from the Neogene under-layer. They could have been redeposited
or people could have used the Neogene sediments as filling for the
stone ramparts.
4.9. Overlays of the cultural layer
Above the cultural layer, humic loam was noticed in the inner
part of the rampart and youngest flood clays, which covered the
destroyed part of the rampart from the outer side. These youngest
flood sediments levelled the flood-plain surface (Opravil,
1983b; Havlícek, 2001). In these samples there was evidence of
the partial regeneration of woodland. The large quantity of Pinus
pollen that is manifest indicates that the process of succession is
still continuing. The same phenomenon is visible in all the
uppermost parts of the profiles and boreholes. This landscape
regeneration seems to be the result of a decrease in the intensity
of human activity.
5. Discussion and interpretations
The occurrence of Carpinus in the sediments dated as Early
Atlantic is noteworthy. Its spreading into the Czech Republic
(Bohemia and Moravia) during the Subboreal is assumed to be
based on a traditional idea (i.e. Firbas, 1949). In addition, Kalis et al.
(2003) does not assume its penetration in the Early Atlantic forests
in Central Europe. Magyari (2002) noted the first appearance of
Carpinus in the SE Carpathians and in the North Hungarian Middle
mountains around 8500 cal BP and their role as an important
element of the woodlands from about 7500 cal BP. Glacial refugia
for Carpinus are referred to in the Balkan Peninsula (Huntley, 1988;
Willis, 1994; Bozilova and Tonkov, 2000) and in restricted areas of
the Hungarian plains (Magyari, 2002; Feurdean, 2005). According
to Ralska-Jasiewiczova et al. (2002), an isopollen map of the
spreading of Carpinus from Italy and Romania shows that it reached
the SE of Poland in 7000 BP. Several authors (Gardner, 2002;
Ralska-Jasiewiczova et al., 2002) presume the expansion of Carpinus
to be an anthropogenic activity (from 6800 cal. BP).
Rybnícková (1985) supposed the much earlier spreading of
Carpinus in the area of the confluence of the Dyje and Morava
rivers than in other regions of the Czech Republic. The scarce
findings of Carpinus in the Mesolithic age and the increased
occurrence of settlements where Linear Pottery was used (Lower
Neolithic) confirm, according to Opravil (1983a), the early arrival
of Carpinus in South Moravia. From the time of the climaticoptimum
Atlantic, macro remains such as wood and nutlets have
been found (Opravil, 1983a, 1984). Our findings of Carpinus (6%)
(Fig. 6) in the layer dated as Mesolithic (V 13 e 8240 cal BP), and
also below it, also confirm the earlier spreading of Carpinus in the
Czech Republic (i.e. South Moravia).
The occurrence of pollen grains of Juglans is very interesting and
disputatious in the area studied. It occurred in nearly all types of
sediment at almost all the depths of this locality (e.g. S3 1.27 m:
7830 Æ 60 cal BP) (Fig. 8). The present area of the natural extent of
Juglans is in the mesophyte forest of the Balkans, northern Turkey,
the Caucasus and Central Asia. The occurrence of Juglans is traditionally
interpreted as an import from southern Europe during
Roman times (Hajnalová, 2001). From several investigations, it is
presumed that there is an older occurrence of it in the northern
Alps. Another example is the coalified Juglans wood that was found
in the Slovakian Neolithic locality Sarisské Michalany (Hajnalová,
2001). Cyprien et al. (2004) presents pollen diagrams from France
(lower Loire) with grains of Cerealia, Fagopyrum and Juglans
occuring at about 6300 cal B.C.; nevertheless, Behre (2007)
recommends a thorough reinvestigation of the site. Griffiths et al.
(2004) assumes Juglans in the pre-Holocene refugias of the Balkans
(Slovenia and Greece). The survival of Juglans during the last
glacial stage is assumed by Carrión and Sánches-Gómez (1992) in
southern Spain. Pollen grains of Juglans were found by Jankovská
et al. (2003) in some handmade boreholes in the neighbouring
Slavonic fort of Mikulcice (flooding sediments of 2.20e2.30 m) and
also by Svobodová (1990) in Pohansko at a depth of 1.60 m. Due to
the presence of redeposited Neogene pollen grains in some Holocene
samples, the eventual redeposition of Juglans pollen could not
be excluded. Long-distance transport by strong winds from
southern areas could be considered.
The findings of Cereal-type pollen in the pre-Neolithic (V
13e8240 cal BP) sediments are disputable. Secondary anthropogenic
indicators P. aviculare, P. lanceolata, R. acetosella, HumulusCannabis
(which could, however, be original to this landscape) were
found. Cereal-type pollen was found in parts of Central Europe
sporadically and known from the pre-Neolithic period, according to
Lang (1994). The first cereal pollen found in the Neolithic (7600 cal
BP) was in south-west Germany (Rösch, 2000). The possibility of
the pre-Neolithic origin of Cereals in Central Europe is discussed by
Behre (2007) and Kalis et al. (2003). According to Zolitschka et al.
(2003), there is a strong influence of human activity on the
natural landscape in the late Neolithic, indicating that agriculture
was expanding over most of Central Europe. These changes coincide
with the start of colder and moister climatic conditions
following the “Holocene climatic optimum” (Davis et al., 2003).
According to Behre (2007) and Beug (2004), apart from cultivated
cereals, several species could belong to the Cerealia-type pollen.
Particularly the Glyceria species, growing in wet habitats, could be
expected in Central European pollen diagrams, according to Behre
(2007). Similar vegetation types are typical for the basal Holocene
pollen spectra in our area of study (Figs. 1, 2 and 4). Long distance
transport of cereal pollen must also be considered (Behre, 2007).
Roszková (2007) found the Triticum-type pollen in the locality at
the Giant Mountains at an altitude of 1471 m a. s. l., which could
have been transported by wind currents from the lowland.
N. Doláková et al. / Journal of Archaeological Science 37 (2010) 2538e25502548
6. Conclusions
Palynological studies were done in the boreholes and profiles
from the inside and outside space of the settlement. The investigated
material was composed of mineral sediment with mostly
a small admixture of organic particles.
The lowermost parts of some boreholes penetrated into the
Upper Miocene sediments of the Vienna basin. The age of the
studied Holocene sediments were dated from the Mesolithic (14
C in
8240 cal BP) up to the 12th century or even more recent times (the
youngest flood loam).
Variations in the proportions of individual plant types of the
studied Holocene layers were probably controlled by the character
of deposition caused by changing humidity as well as by human
activity (clearance, agriculture, pasture). The predominance of
flood-plain forest against the mesophyte oak-lime-hornbeam
forest, which is visible (several times) in the pollen spectra, indicates
increased aerial and edaphic humidity. The first significant
decrease of arboreal pollen was detected in the level preceding the
lower flood loams and lower soil horizon. There is evidence of
human impact in these layers (the occurrence of Cerealia, field
weed such as P. aviculare, C. cyanus, as well as secondary anthropogenic
elements P. lanceolata, R. acetosella, Humulus e Cannabis
and Chenopodiaceae).
These facts indicate that there was agriculture in the Neolithic
period e in the time before the sedimentation of the flooding loam
in the bottom land. There may have been human activity at the first
accumulation of these flooding sediments.
A striking human influence was also registered in the layers
dated as Hallstatt.
The partial rejuvenation of forest was visible in the overlaying
horizons. This rejuvenation was followed by the greatest deforestation
in the Great Moravian cultural horizon.
The findings of pre-Neolithic Cereal-type pollen such as Juglans
are disputable.
The palynomorphs demonstrate that the stone ramparts served
not only as protection against invasions by Hungarians, but additionally
as a flood prevention barrier.
The occurrence of Carpinus pollen in the layer dated as Early
Atlantic, and also beneath it, confirms the earlier spreading of
Carpinus in the Czech Republic (i.e. South Moravia). This concurs
with earlier findings of macro remains from the old settlement
regions of the Czech Republic.
Acknowledgements
The study was supported by Research project MSM0021622427
(Czech Republic) Inter-disciplinary Centre for Research into
Prehistoric to High Medieval Social Structures.
We are grateful to Sarka Rousava for correction of the English
language and to anonymous reviewers for constructive comments.
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63
SBORNÍK NÁRODNÍHO MUZEA V PRAZE AC TA M U S E I N AT I O N A L I S P R AG A E
Řada B – Přírodní vědy • sv. 67 • 2011 • čís. 1–2 • s. 63–71 Series B – Historia Naturalis • vol. 67 • 2011 • no. 1–2 • pp. 63–71
BADENIAN (LANGHIAN – EARLY SERRAVALLIAN) PALYNOFLORA FROM THE CARPATHIAN
FOREDEEP AND VIENNA BASIN (CZECH AND SLOVAK REPUBLICS)
NELA DOLÁKOVÁ
Masaryk University,Faculty of Sciences,Institute of Geological Sciences,Kotlářská 2,611 37,Brno,Czech Republic,
e-mail:nela@sci.muni.cz
MARIANNA KOVÁČOVÁ
Comenius Universtity,Faculty of Sciences,Department of Geology and Palaeontology,Mlynská dolina G,SK-84215 Bratislava,Slovak
Republic,e-mail:kovacova@fns.uniba.sk
PETRA BASISTOVÁ
Masaryk University,Faculty of Sciences,Institute of Geological Sciences,Kotlářská 2,611 37,Brno,Czech Republic,
e-mail:162649@sci.muni.cz
Doláková, N., Kováčová, M., Basistová, P. (2011): Badenian (Langhian – Early Serravallian) palynoflora from the Carpathian Foredeep
and Vienna Basin (Czech and Slovak Republics). – Acta. Mus. Nat. Pragae, Ser. B, Hist. Nat., 67(1-2): 63-71, Praha.
Abstract. The Badenian (Langhian - Early Serravallian) marine sediments from the adjacent areas within the Central Paratethys and NE
part of the Vienna Basin, were studied from a palynological perspective. The pollen data document a subtropical climate during the Miocene
Climatic Optimum with dominant representation of zonal vegetation being evergreen broadleaved forests. Higher differentiation of
oak type pollen, increasing number of Platanus pollen and different types of herb were observed.
Some thermophilous elements (especially Sapotaceae, Palmae, Mastixiaceae and Lygodiaceae) decreased and there was an increase
of the warm to cold temperate zone taxa which were first registered during the Late Badenian. These findings together with a higher proportion
of extrazonal vegetation (Tsuga, Picea and Abies) in the Late Badenian pollen spectra document changes due to the uplift of the
Carpathian Mountain chain.
n Palynology, Badenian, Langhian, Early Serravallian, Carpathian Foredeep, Vienna Basin
Received December 10, 2010
Issued August 2011
Introduction
The adjacent areas within the Central Paratethys in the Czech
and Slovak republics, the Carpathian Foredeep and Vienna
Basin, were studied from a palynological perspective. The
Lower Badenian (early Langhian) transgressions from the
Mediterranean toward the Central Paratethys realm flooded
the Pannonian Basin and continued along straits in the
Carpathian Chain into the Carpathian Foredeep. During the
Lower Badenian, in addition to the shallow partial basins,
depressions of unstratified calcareaous clay developed which
were more than a hundred metres deep – "tegels" (Brzobohatý
1982, 1997; Chlupáč et al. 2002). The isolation of eastern
parts of the Central Paratethys at the end of this period
(Late Langhian) resulted in the “Middle Badenian” salinity
crisis. Thick evaporite sediments were deposited in the Carpathian
Foredeep. During the Upper Badenian (Early Serravallian),
the most recent total marine flooding covered the
whole back-arc basin and a great part of the foredeep
(Kováč et al. 2007). The globally observed Middle Miocene
Climatic Optimum is, according to Böhme (2003) and
Utescher et al. (2000), clearly reflected in the studied Badenian
material. Mountain chain uplift and strong relief
development is documented by the origin of river drainage
feeding into the huge deltaic systems of the back arc basin
(Kováč 2000; Konečný et al. 2002).
The lowermost Badenian strata, which can be recognized
almost everywhere in the Central Paratethys realm,
contain planktonic foraminiferal assemblages in which the
genus Praeorbulina is associated with the genus Orbulina
in the calcareous nannoplankton Sphenolithus heteromorphus
Zone NN5 (Berggren et al. 1995; Fornaciari, Rio
1996). The time span of the Late Badenian is approximately
coeval to the upper part of the MN7 Globorotalia peripheroacuta
Lineage Zone of Berggren et al. (1995) and the lower
part of the Discoaster exilis Zone NN6 (Martini 1971).
Material and methods
In total 64 Lower Badenian and 39 Upper Badenian
samples of marine clays were studied palynologically.
The Lower Badenian samples came from the boreholes
Ivaň, Rebešovice, Chrlice, Opatovice, Otmarov, Přemyslovice
and outcrops Brno-Královo Pole, Moravské Knínice
64
and Sivice, situated in the southern part of the Carpathian
Foredeep and from the three regional stratotype localities
Oslavany (OV-1), Židlochovice (Ž–1) and (Ž–2). The Late
Badenian sediments came from the boreholes Gajary 23,
Sekule 1, Jakubov 54, Zohor 1, Lozorno and outcrop at the
faciostratotype locality Devínska Nová Ves situated in the
Vienna Basin.
Standard maceration with HCl (20%), HF, KOH and
HCl (10%) was carried out. Due to the increasing number
of palynomorphs, heavy liquid (ZnCl2) with a density of
2g/cm3 was utilised. Pure glycerine or glycerine gelatine
were most frequently used as the observation media.
The percentage of the individual taxa were calculated
from the total sum of a minimum 150 determined pollen
grains and spores.
The palaeotropical and arctotertiary elements are classified
based on the Neogene pollen flora of Central Europe
(Stuchlik et al.1994). The vegetation units terminology was
used according to Kvaček et al. (2006) and Kovar-Eder et
al. (2008).
To resolve the situation with problematic taxa identification,
Quercus, Platanus, selected herbs, Castanea x Castanopsis,
a recent Castanea pollen material has been studied
under SEM. These observations and photos were done
using Scanning Electron Microscope JEOL JSM – 649 OLV
at the Institute of Geological Sciences, Masaryk University
in Brno.
Vegetation
The Lower Badenian palynospectra were rich in Dinoflagellata
and foraminiferal linings (Pl. 1, figs 1-3). Sporadic
occurrences of Botryococcus (Pl. 1, fig. 4) and pollen
of aquatic coastal plants Sparganium, Potamogeton and
Utricularia (Pl. 2, fig. 25) indicated a fresh water influence
on some facies.
The proportion of zonal vegetation with evergreen broadleaved
forests (Sapotaceae, Engelhardia, Platycarya, evergreen
Fagaceae – Castanopsis, Trigonobalanopsis, morphotypes
Tricolporopollenites henrici, T. microhenrici, T. liblarensis,
Reevesia, Cornus-Mastixia, Rutaceae and Araliaceae
and Pteridaceae) represents up to 30% of the pollen
spectra (Pl. 2). In the Miocene time interval, the thermophilous
morphotaxa Gothanipollenites gothani and
Clerodendrumpollenites microechinatus were first found in
Lower Badenian sediments of the studied area.
The broad-leaved deciduous elements of warm – temperate
mixed mesophytic forests such as Quercus, Castanea,
Carya, Celtis, Juglans, Tilia, Zelkova, Ostrya, Carpinus,
Betula and Cercidiphyllum generally did not exceed
10%. A higher diversity of ”oak type” pollen grains, e.g.
Quercus robur-pubecscens, were recorded in the pollen
spectra (Pl. 2, figs 6-10, Pl. 4, figs 7-12). Cercidiphyllum
and Castanea/Castanopsis were identified from the Lower
Badenian pollen spectra (Pl. 2, fig. 15; Pl. 3, figs 1-15).
The azonal vegetation was represented by riparian forests
with Ulmus, Alnus, Fraxinus, Liquidambar, Salix and
Lythraceae and the coastal swamps by Nyssa, Sciadopitys,
Taxodiaceae, Cyrillaceae and Myricaceae. Pollen grains of
Platanus ipelensis sensu Pacltová (1984) were abundantly
present for the first time in the Lower Badenian taphocenoses
(Pl. 2, figs 11-14, Pl. 4, figs 10-12). In the pollen
spectra herbs and heliophilous elements Poaceae, Asteraceae,
Caryophyllaceae, Chenopodiaceae, Ericaceae and
Ephedra were regularly present, less frequent were Urtica,
Plantago, Salvinia and Lavandula (Pl. 2). There were
noticeable polyporate pollen grains with microechinate perforate
sculptures visible under SEM, they were determined
as Caryophyllaceae cf. Saponaria (Pl. 4, figs 1-3), or without
perforations – Thalictrum type (Pl. 4, figs 4-6).
An extremely high proportion (more than 60%) of Pinaceae
pollen was present in some samples (Pl. 1). From the
borehole Oslavany and the uppermost parts of boreholes
Židlochovice 1, practically only Pinaceae and dinoflagellates
were found. This could be due to taphonomical and
ecological reasons (long air-transport range and therefore
accumulation in marine sediments distant from the sea-
shore).
In comparison with the Lower Miocene spectra, portion
of the thermophilous elements P1 sensu Stuchlik et al.
(1994), Symplocos, Sapotaceae, Palmae, Mastixiaceae and
Lygodiaceae started to decrease since the Badenian (Doláková,
Slamková 2003; Doláková et al. 1999). An increased
proportion of the arctotertiary taxa (Quercus, Ulmus and
Carya) were noted in the Upper Badenian palynospectra
from the Vienna Basin. Thermophilous elements (Platycarya,
Engelhardia, Myrica, Distylium and thermophilous
Fagaceae) were still present, but Sapotaceae had disappeared.
Herbs were represented predominantly by the halophytic
taxa – mainly Chenopodiaceae. The higher proportions
(up to 30%) of extrazonal (mountain) vegetation in the
pollen spectra (Picea, Abies, Tsuga, Cedrus) were first
recorded from the Upper Badenian. The main reason of this
phenomenon is the uplift of the Carpathian Mountain chain
and subsidence of adjacent lowlands.
Discussion
Lower Badenian macrofloristic remains from the
Carpathian Foredeep are rare. Only in the Lower Badenian
marine sandstones at Smolín near Pohořelice, Knobloch
(1963, 1968) and Knobloch et al. (1975), described poorly
preserved Lauraceae leaves Daphnogene bilinica, and Betulaceae
leaves. The macrofloristic taphocenoses from the
vicinity of Česká Třebová were described by Knobloch
(1968); Knobloch et al. (1975) as an association totally
dominated by arctotertiary elements (prevailing being Myrica
lignitum, Alnus cf. feroniae, Pinus, Salix, Populus, Fagus,
Pterocarya, Parrotia, Ulmus and Fraxinus).
Equally previously published palynological results from
the Lower Badenian sediments of the Carpathian Foredeep
(Basistová, 2009; Bruch et al. 2004; Hladilová et al. 1999,
2001) our results indicate a more thermophilous – subtropical
character of the climate.
Macrofloristic findings from the Upper Badenian were
described at the localities Opava – Kateřinky, and borehole
Smolkov near Opava (Knobloch 1968). An absence of laurophylous
leaves was observed and a generally warm – temperate
character of the paleoclimate equaly to sediments
with Lower Badenian floras.
In the Upper Badenian pollen spectra from boreholes
OS-1 Kravaře and OS-2 Hať in the vicinity of Opava, the
arctotertiary elements (with a high proportion of Pinaceae,
65
partly mountain vegetation) dominated and thermophilous
taxa were represented only by Engelhardia and Lygodiaceae
(Cicha et al. 1985).
Planderová and Gabrielová (1975) noticed a decrease in
the most thermophilous elements (Sapotaceae, Symplocos,
Palmae, Mastixiaceae and Lygodiaceae) in the Lower Badenian
pollen spectra in comparison with Early Miocene
spectra. Based on this data they interpreted a subtropical
climate with humidity oscillations during the Badenian time
interval. Based on the Upper Badenian pollen spectra Planderová
(1990) interpreted a drier and colder climate in comparison
with the Lower Badenian time span. In comparison
with the Karpatian pollen spectra, the proportion of herbs
and heliophillous elements in Badenian material is higher
(Doláková and Slamková 2003).
Holcová et al. (1996) observed up to 30% thermophilous
taxa in the Lower Badenian pollen spectra from the
borehole N-95 in Strháry-Trenč graben from the South Slovakia
basin.
Based on many palynological studies, Oszast and Stuchlik
(1977), Łancucka-Srodoniowa (1966) and Dyjor and
Sadowska (1984) observed some differences between Badenian
floras from the Lowlands in Southern Poland (more
subtropical and warm – temperate) and from the mountain
regions (2-3 altitudinal zones with warm-temperate mixed
forests and with coniferous forests in the upper parts).
Planderová et al. (1993a,b) noted a lack of paleotropical
elements of the P1 group in the Lower Badenian in the
northern part of the Paratethys area. In the Upper Badenian
they observed these elements only in the southern Paratethys
area (Hungary and former Yugoslavia). Based on the
flora of Parschlung (Karpatian/Early Badenian) Kovar-Eder
et al. (2004) indicated a drier warm-temperate climate with
relatively rare subtropical humid elements, while subhumid
sclerophyllous woody taxa were well represented.
Kováč et al. (2008) and Kvaček et al. (2006) previously
published palynological studies of Badenian sediments. In
comparison with these data it is evident that even though
the Badenian climate was generally subtropical, the proportion
of key termophilous taxa rapidly decreased in the
Upper Badenian.
Conclusions
Pollen flora from the Lower Badenian sediments indicated
the presence of evergreen broadleaved forests, constituting
up to 30% of the pollen spectra. In comparison with
the Lower Miocene time span the proportion of thermophilous
elements, Sapotaceae, Symplocos, Palmae, Mastixiaceae
and Lygodiaceae, decreased in the Badenian pollen
spectra. A greater differentiation of “oak type” pollen and
a higher amount of Platanus pollen were recorded in the
studied samples which corresponds with the results of
Knobloch and Kvaček (1996).
Herbs and heliophilous vegetation started to be more
frequent than in Lower Miocene floras.
In comparison with macrofloristic findings, the pollen
grains indicate that the Lower Badenian floras had a more
thermophilous character. Due to the need for more detailed
analyses and correlation, this study will continue on a larger
scale and Badenian sediments from Hungary, Slovenia,
and Austria will be analysed in the future.
Initiation of altitudinal zonation (Kováč et al. 1998, Kováčová
et al. in press) is documented by an increase of
mountain elements and arctotertiary taxa (Quercus, Ulmus
and Carya), a decrease of thermophilous elements (Platycarya,
Engelhardia, Myrica, Distylium and thermophilous
Fagaceae), and the disappearance of Sapotaceae in the palynospectra
during the Upper Badenian.
Acknowledgements
The study was supported by the Projects 205/09/0103
(Grant Agency of the Czech Republic), APVV-0280-07,
VEGA 2/0060/09, ESF-EC-009-07, VEGA 1/0483/10.
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Explanation of the plates
PLATE 1
1. Foraminiferal tapetum, Židlochovice 1 (2.3 m).
2. Marine dinoflagellates, Židlochovice 2 (8.9 m).
3. Marine dinoflagellates: a – LM, b – SEM, Židlochovice
1 (11.1 m).
4. Botryococcus sp., Židlochovice 1 (7 m).
5. Cathayapollis potoniei (SIVAK) ZIEMBIŃSKA-TWORZYDŁO,
Židlochovice 1 (7 m).
6. Cedripites miocaenicus KRUTZSCH, Židlochovice 1 (8,5 m).
7. Cathayapollis sp., SEM, a – whole pollen grain, b – detail,
Židlochovice 1 (11.1 m).
PLATE 2
1. Sapotaceae: Sapotaceoipollenites sapotoides (PFLUG et
THOMSON) POTONIÉ; Moravské Knínice.
2. Mastixioideae: Cornaceaepollis satzvayensis (PFLUG)
ZIEMBIŃSKA-TWORZYDŁO, Židlochovice 1 (10.9 m).
3. Quercoidites henrici (POTONIÉ) POTONIÉ, THOMSON et
THIERGART, Ivaň (43.75 m).
4. Platycaryapollenites miocaenicus NAGY, Židlochovice
1 (10.9 m).
5. Hedera type: Araliaceoipollenites reticuloides THIELEPFEIFER;
Židlochovice 1 (9.7 m).
6.–10. Quercus robur-pubescens type: Quercoidites
granulatus (NAGY) SLODKOWSKA; 6,7 – Židlochovice
(excavation); 8 – Židlochovice 1 (10.3 m); 9,10 – Moravské
Knínice.
11.–14. Platanus sp.: Platanipollis ipelensis (PACLTOVÁ)
Grabowska; 11,12 – Židlochovice 1 (10.3 m); 13 – Židlochovice
1 (11.1 m).
15. Cercidiphyllum sp.: Cercidiphyllites minimireticulatus (Trevisan)
ZIEMBIŃSKA-TWORZYDŁO, Židlochovice 1 (10.9 m).
16. Urtica sp.: Triporopollenites urticoides NAGY, Židlochovice
2 (16 m).
17. Plantago sp.: Plantaginacearumpollenites miocaenicus
NAGY, Židlochovice 1 (10.3 m).
18. Lavandula sp.: a,b, Židlochovice 1 (0.9 m).
19. Salvia verticillata type; Židlochovice 1 (9.7 m).
20.–23. Caryophyllaceae gen. indet.: Caryophyllidites microreticulatus
NAGY, 20 – Židlochovice 1 (10.9 m); 21,
22 – Židlochovice 1 (7 m); 23 – Židlochovice 1 (10.3 m).
24. Galium type, Židlochovice 1 (9.7 m).
25. Utricularia sp.; Židlochovice 1 (9.7 m).
26. Asteraceae: Tubulifloridites macroechinatus (Trevisan)
NAGY, Ivaň (16.2 m).
27. Asteraceae: Cichoreacidites gracilis NAGY, Ivaň 43.7 m).
PLATE 3
1.–9. Castanea sp. – recent pollen, 1–6 LM; 7–9 SEM.
10.–12. Castanea sp.: Tricolporopollenites cingulum (POTONIÉ)
oviformis THOMSON et PFLUG – form A,
Židlochovice1 (11.1 m), 10 – LM; 11–12 – SEM.
13.–15. Castanopsis sp.: Tricolporopollenites cingulum
(POTONIÉ) oviformis THOMSON et PFLUG – form B,
Židlochovice 1 (11.1 m), 13 LM; 14–15 SEM.
16.–18. ?Fagaceae: Tricolporopollenites liblarensis (THOMSON)
Grabowska, Židlochovice 1 (11.1 m), 16 –
LM; 17–18 – SEM.
PLATE 4
1.–3. Caryophyllaceae: cf. Saponaria sp., Židlochovice
1 (11.1 m), 1 – LM; 2–3 – SEM.
4. –6. cf. Thalictrum sp., Židlochovice 1 (11.1 m), 4 – LM;
5–6 SEM.
7. –9. Quercus sp., Židlochovice 1 (11.1 m), 7 – LM; 8–9
SEM.
10.–12. Quercus sp., Židlochovice 1 (11.1 m), 10 – LM;
11–12 SEM.
13.–15. Platanus sp.: Platanipollis ipelensis (PACLTOVÁ)
Grabowska, Židlochovice (11.1 m), 13 – LM;
14–15 – SEM.
LM – light microscope
SEM– scanning electron microscope
Magnification of all photographs is indicated directly in
figures.
68
PLATE 1
69
PLATE 2
70
PLATE 3
71
PLATE 4
1 3
Facies (2014) 61:419
DOI 10.1007/s10347-014-0419-z
ORIGINAL ARTICLE
The Langhian (Middle Badenian) carbonate production event
in the Moravian part of the Carpathian Foredeep
(Central Paratethys): a multiproxy record
Katarína Holcová · Juraj Hrabovský ·
Slavomír Nehyba · Šárka Hladilová · Nela Doláková ·
Atilla Demény 
Received: 30 January 2014 / Accepted: 1 September 2014
© Springer-Verlag Berlin Heidelberg 2014
carbonates were transported to the outer shelf by gravity
flows. Climatic instability and relative sea-level changes,
induced mainly by substantial tectonic activity, caused the
carbonate bodies to be small with a high ratio of siliciclastic
components, indicating only a short-term and spatially
restricted environment suitable for carbonate production.
Exceptionally, carbonate production persisted longer during
the whole sea-level cycle (“Rousínov Ridge”). Siliciclastic
intercalations in these larger limestone bodies represent
catastrophic rain events that transported a higher
amount of terrigenous material into the basin. The specific
climatic conditions of the carbonate production event,
namely climatic instability and aridification with episodic
intensive rain, were associated with the Middle Miocene
climatic transition in the study area.
Keywords  Carbonate–siliciclastic complex ·
Paleoecology · Middle Miocene climatic transition ·
Langhian · Carpathian Foredeep
Introduction
The study area, the Central Paratethys, represents a chain of
Oligocene and Miocene epeiric seas in Central and Eastern
Europe with marked oscillations of paleoecological parameters
and episodic communication with the oceanic realms
(Rögl 1999). Siliciclastic sedimentation strongly prevailed
in the basin. Rare occurrences of carbonates were recorded
from the Early Miocene (Nebelsick 1989), but only carbonates
from the Middle Miocene have been described from
many places. In addition to the studied Moravian part of the
Carpathian Foredeep (Doláková et al. 2008), they were also
recorded in the Eastern Alpine Foredeep (Mandic 2004),
the Vienna and Styrian basins (Schaleková 1973; Baráth
Abstract  The carbonate production event in the Moravian
part of the Carpathian Foredeep is known as a deposition
of a carbonate–siliciclastic complex in the marginal
part of the basin, correlating with the time period from
the last occurrence of Helicosphaera waltrans (14.36 Ma)
to the last occurrence of Sphenolithus heteromorphus
(13.34 Ma). Sedimentological and microfacial data, analysis
of foraminifera, calcareous nannoplankton, red algae,
mollusks, palynology, as well as oxygen and carbon stable
isotopes from foraminiferal tests, were used to interpret
the specific paleoenvironment of the carbonate production
event. The event was accelerated by a decrease of terrigenous
input due to a large transgression and, primarily,
an increasingly arid climate. Production of carbonate was
related to oligotrophic conditions, expansion of sea-grass
meadows, summer downwelling circulations and winter
stratification of the water column. Autochthonous and
semi-autochthonous carbonates were deposited in shallow-water
near the fair-weather wave-base; allochthonous
K. Holcová (*) 
Institute of Geology and Paleontology, Charles University
in Prague, Albertov 6, 12843 Prague 2, Czech Republic
e-mail: holcova@natur.cuni.cz
J. Hrabovský · S. Nehyba · N. Doláková 
Departure of Geology and Paleontology, Masaryk University,
Kotlárˇská 2, 600 00 Brno, Czech Republic
Š. Hladilová 
Department of Biology, Faculty of Education, Palacký University,
Purkrabská 2, 771 40 Olomouc, Czech Republic
A. Demény 
Institute for Geological and Geochemical Research,
RCAES, Hungarian Academy of Sciences, Budaorsi ut 45,
Budapest 1112, Hungary
	 Facies (2014) 61:419
1 3
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et al. 1994; Kysela 1988; Riegl and Piller 2000; Wiedl et al.
2012), the Pannonian Basin in Hungary (Randazzo et al.
1999; Moissette et al. 2007) and Slovakia (Vass et al. 2007),
the Zrin-Dvor Basin in Croatia (Basso et al. 2008; Martinuš
et al. 2012) and Bosnia–Herzegovina (Pezelj et al. 2013), the
Carpathian Foredeep in Poland (Pouyet and Tarkowski 1998)
and Ukraine (Radwan´ski et al. 2006) and the Transylvanian
Basin in Romania (Filipescu and Gîrbacea 1997).
The aim of this study is to evaluate the influence of
global and local paleoclimatic, paleogeographic and tectonic
factors on carbonate production in the Moravian part
of the Carpathian Foredeep based on a multiproxy analysis.
Geological setting
The Early–Middle Miocene Carpathian Foredeep is a peripheral
foreland basin that developed from subsurface loading
of the Alpine-Carpathian orogenic belt on the passive margin
of the Bohemian Massif. The Carpathian Foredeep exhibits
striking lateral variations in basin width, depth and stratigraphy
of deposits, along with variations in the pre-Neogene
basement composition and tectonic subsidence. The basin
continued to the south (the Alpine Foredeep/Alpine Molasse
Zone) and to the northeast (the Polish part of the Carpathian
Foredeep) (Oszczypko 1998; Nehyba et al. 2008a).
The distinctive geometry of the Early Badenian deposits
reflects the important reconstruction of the basin. The location
of the basin and the distribution and character of the
deposits could be explained by the tectonic development of
the Carpathian orogenic wedge. Both eustasy and tectonics
governed the depositional architecture of the basin during
the Early–Middle Badenian (Nehyba and Šikula 2007;
Nehyba et al. 2008a).
Pelitic sediments (“Tegel”) with sandstone intercalations
and biohermal bodies strongly dominate volumetrically
within the basin with a maximum thickness of
about 600 m. These mudstones vary in silt and clay content,
amount of shell debris, intensity of bioturbation and
sedimentary structures. They were interpreted as predominantly
outer shelf deposits or hemipelagites (Papp et al.
1978; Cicha 2001; Nehyba et al. 2008a). Limestone bodies
are represented by lenses in siliciclastic complexes. Their
thickness varies by a few meters, occassionally reaching
a few tens of meters (maximum 40 m). Limestone bodies
(called “Leitha-limestones” or “Lithothamnian-limestones;
Papp et al. 1978) are concentrated in three areas and their
positions in the succession are not necessarily isochronous.
Their occurrences are connected with the prograding
coast line in the western and north-western margins of the
preserved Carpathian Foredeep in Moravia. At the eastern
margin, redeposition of the red-algal limestones into more
internal parts of the basin apparently played an important
role (Doláková et al. 2008). Biohermal bodies, restricted
both areally and in thickness, are represented mainly by red
algal limestones and, more rarely, by bryozoan bioherms.
The rich fossil content of the bioherms has attracted the
attention of paleontologists since the nineteenth century;
many works describing individual fossil groups originate
from the 2nd half of the 20th century. These works are summarized
by Hladilová and Zdražílková (1989) and Doláková
et al. (2008). Most recent contributions were focused
on the bryozoan limestones from Podbrˇežice (Zágoršek and
Holcová 2005: Bryozoa and Foraminifera; Nehyba et al.
2008b: oxygen and carbon isotopes from bryozoan fragments;
Hrabovský et al. 2015). However, a detailed multiproxy
analysis of the ecosystem evolution is missing.
Material and methods
This work is focused on a detailed analysis of two limestone
bodies north of the town of Rousínov in the middle
part of the Carpathian Foredeep in Moravia (Czech
Republic): the sections of Kroužek (molluscan and bryozoan
biodetritic and biomicritic limestone) and Podbrˇežice
(bryozoan biodetritic limestone/bryozoan bank) (Hladilová
and Zdražílková 1989; Fig. 1). These sections, situated on
the western margin of the Foredeep basin, were compared
with limestone bodies from boreholes in Židlochovice situated
at the eastern margin of this marine basin (Doláková
et al. 2014) and with denudation relicts in Middle Badenian
deposits (including carbonates) from Kralice (Zágoršek
et al. 2007, 2009).
Microfacies and red algae were studied in thin-sections.
The ratio of microfacial components was evaluated using
JMicrovision software and the point-counting method.
They were evaluated according to a 250–400 points/thinsection
scale. Quantitative data were collected from 24
thin-sections. Twelve components were distinguished in
the samples (Table 1). Components were recognized and
classified according to Flügel (2004). We classified mixed
siliciclastic limestones with more than 10 % of lithoclasts
according to Mount (1985). Limestones with less than
10 % of lithoclasts were classified according to Dunham
(1962) and Wright (1992). Woelkerling (1988) and Braga
et al. (1993) were followed in generic description of coralline
algae (Sporolithalles, Corallinales, and Rhodophyta).
Foraminifera and molluscs were studied from 63–
2,000 μm fractions. Molluscs were studied from 38 samples
(Kralice nad Oslavou: 12 samples; Židlochovice boreholes:
ŽIDL1—13 samples, ŽIDL2—8 samples) and foraminifera
from 106 samples (Podbrˇežice: 21 samples; Kroužek: 24
samples; Kralice nad Oslavou: 12 samples; Židlochovice
boreholes: ŽIDL1—26 samples and ŽIDL2—23 samples).
Ultrasonic treatment was used for further cleaning of
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Fig. 1  Locations of sections studied (a) and their lithology (b)
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molluscs. For the molluscan fauna from Kroužek, the material
of Šob (1940), kept in the collections of the Moravian
Museum in Brno (Hladilová 1984), was studied. About 200–
300 specimens of foraminifera from each sample were determined
and relative abundances of taxa, as well as plankton/
benthos ratios (P/B ratio), were calculated.
Benthic foraminiferal assemblages and calcareous nanoplankton
were statistically classified using the multivariate
techniques of PAST software (Hammer et al. 2001).
Calcareous nannoplankton was studied from the same
106 samples as the foraminifera. The abundance of nannoplankton
was expressed semi-quantitatively as the number
of specimens in the visual field of the microscope (Zágoršek
et al. 2007). About 200–500 specimens of calcareous nannoplankton
were determined from individual samples and
the relative abundances of taxa were calculated.
For palynological studies, two samples from sandy siltstones,
one limestone sample of the Podbrˇežice section
and four samples from sandy siltstones of the Kralice section
were studied (Fig. 1). The samples were treated with
cold HCl (35 %) and HF (70 %), removing carbonates and
silica. Separation of the palynomorphs from the rest of the
residue was carried out using ZnCl2 (density = 2 g/cm3
).
No Tertiary palynomorphs were observed in the limestone,
the only samples containing contemporary pollen
contamination. Palynomorphs were recorded only in siliciclastic
intercalations in carbonates from the Židlochovice
boreholes (Doláková et al. 2014).
Oxygen and carbon isotopic composition tests of the
foraminiferal can be evaluated only for well-preserved
samples from the Židlochovice and Kralice sections. The
methods used were described by Holcová and Demeny
(2012). Groups of small-sized, four-chambered Globige‑
rina sp. and Cibicidoides spp. were chosen for isotopic
analysis. Planktonic foraminifera for isotopic analysis
were picked from a size fraction of 63–200 μm, in which
no specimens with reproductive chambers were observed.
Cibicidoides spp. were picked from the 63–300 μm fraction.
The carbon and oxygen isotope compositions of calcite
were determined at the Institute for Geological and
Geochemical Research, RCAES, Hungarian Academy of
Sciences (Budapest, Hungary).
Results
Biostratigraphy
Biostratigraphic correlation of the carbonate production
event was based on the occurrence/absence of planktonic
Table 1  Volume of identified
components in the study
samples, counted using
JMicrovision
The code of each thin-section is
in the left column and identified
microfacies (MicF) are in far
right column
CRA coralline red algae, BRY
bryozoans, Mol mollusks, FRM
foraminifera, serpulids, SRP
polychaetes, BRC brachiopods,
UNS unidentified allochems,
PRS pores, LIT lithoclasts,
MIC micrite, SPR sparite
CRA BRY MOLL FRM ECHN SRP BRC UNA PRS LIT MIC SPR MicF
27611151 17.4 17.2 2.7 0.3 0.3 1.2 2 14.3 4.9 4.9 34.2 0.3 3
27611251 10.7 32.5 3.5 0.2 0.2 2.1 1.4 2.3 10.7 1.9 28.3 6.3 3
27611351 1.5 41.6 0.5 1 0.3 0.8 0 2.3 14.8 2.5 26.6 8.3 3
276111251 10.7 59.9 2.7 0 0.3 0 0 0.3 7.7 1 16.2 1.3 3
27611551 52.5 16.2 0.3 0.8 0 0.3 0 3.7 6.7 2 16.2 1.5 3
27611451 27.3 35.3 1 2.3 0.5 0 0 3.3 2.8 3.3 18.8 6.3 3
27611651 20 31.3 1 1.5 0.3 0.3 0 1.5 1.3 4.3 32.5 6.3 3
27611751 7 43 0.7 1.7 0.2 0.2 0 2.9 0.5 1.2 40.5 1.9 3
27611851 4.2 43.8 1 2 0.3 0.8 0.8 6.2 0.5 2.5 33.1 5 3
276111351 5.2 41.3 0.3 1 0 5.2 0 5.2 5.5 2.2 28.9 5.2 3
27611951 6.2 44.6 1.8 0.8 0 0.3 0 5 0.5 3 22.4 15.5 3
276111051 6.7 35.7 12.2 1.5 0 0 0 5.5 1.5 1 25.7 10.2 3
276111151 3.7 37.9 2.2 1 0.5 0 0 6 1.8 4 37.7 5.2 3
17311351 26.5 16.8 17.3 2.2 0 0.7 0 1.5 4.5 8.2 4.5 17.8 1
17311451 17.9 24.6 5.5 2.7 0 0.3 0 5.2 4 10 5.5 24.4 1
17311751 53.9 11 3.7 1.3 0 1 0 2.7 1 4 4.5 18 1
17311951 40.3 15.3 8 0 0 1.8 0 4 7.5 3.5 6.5 13.3 1
173111051 25.2 18 10.7 0.74 0.3 0.5 0 3.2 8.2 7.9 1 24 1
173111151 16.2 15.5 92 1.25 2.5 0 0 4 5.5 16.5 1.5 31.4 1
17311251 6.2 40.1 3.2 2.5 0.8 0.3 0 6.7 3.2 12.4 14.4 10.2 2
173111251 6.5 33.9 2.5 2.7 0 2.5 0 5 6.7 13.2 14.5 12.5 2
173111351 10.7 23.6 7.5 1.5 0 0 0 4.5 17.2 9.2 6.2 19.7 2
17311551 4.5 18.9 2 5.5 0 0.8 0 16.7 1.7 15.9 12.2 21.9 2
17311651 4.3 24 3 2.3 0 1 0 13 1.8 16.8 16.8 17.3 2
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foraminifera and calcareous nannoplankton index species.
The dating of the first and last occurrences (FO, LO)
of index species differs in the world’s oceans (Gradstein
et al. 2012) and in the Mediterranean area (Di Stefano et al.
2008; Abdul Azis et al. 2008; Hüsing et al. 2010). The differences
are summarized in Fig. 2. Due to the connection
of the Central Paratethyan Sea with the Mediterranean, the
correlations of bioevents with the Mediterranean dating are
more probable. Our succession of bioevents follows well
the successions in the Mediterranean area (Di Stefano et al.
2008; Abdul Azis et al. 2008; Hüsing et al. 2010), where
the LO of Praeorbulina spp. is an indefinable event that
may be observed above/before the LO of Helicosphaera
waltrans. The same succession of bioevents was described
in the Carpathian Foredeep by Švábenická (2002).
Of the index species, Praeorbulina (LO: 14.9 Ma; Gradstein
et al. 2012; Mediterranean area: Abdul Azis et al.
2008), Orbulina (FO: 15.1 Ma; Gradstein et al. 2012;
Mediterranean: 14.6 Ma; Abdul Azis et al. 2008; Hüsing
et al. 2010) and Sphenolithus heteromorphus (LO: 13.53
Ma; Gradstein et al. 2012; Mediterranean area: 13.419 Ma;
Abdul Azis et al. 2008; Hüsing et al. 2010) were recorded
in all rocks included in the carbonate production event.
Helicosphaera ampliaperta (LO: 14.91 Ma; Gradstein et al.
2012) and H. waltrans [last common occurrence (LCO) in
the Mediterranean area 14.357 Ma; Abdul Aziz et al. 2008]
are missing.
According to the above-mentioned data, the carbonate
production event can be correlated with the upper part of
the NN5 Zone of calcareous nannoplankton and the M6
Zone of planktonic foraminifera (Berggren et al. 1995);
numerically, the range from 14.36 Ma (LO of H. waltrans)
to 13.34 Ma (LO of S. heteromorphus) can be estimated
based on the Mediterranean dating of bioevents. In the local
Central Paratethys stratigraphy, this interval represents the
Middle Badenian (Hohenegger et al. 2014).
Lithology
The mixed siliciclastic-carbonate complex can be lithologically
characterized by alternations of the following:
(1) Mudstone lithofacies revealing dominant deposition
from suspension in relatively calm conditions. Variations
in the sand content, bioturbation, preservation of planar
bedding and shell debris also reflect periods of a relatively
higher input of material transported by currents in traction
(storms?); (2) Sandy lithofacies and their alternations
with mudstone lithofacies can be connected with deposition
in the lower shoreface or transition zone to a deeper
environment. The sharp bases of the beds, occurrence of
transported limestone clasts, shell debris and planar lamination
suggest a higher flow regime. The absence of clear
wavy structures points to deposition below the fair-weather
wave-base; (3) Heterolithic facies support rapid fluctuation
of siliciclastic input into the environment. Limestone beds,
10–110 cm in thickness, indicate stable conditions of deposition
and severe reduction of siliciclastic input. Multiple
alternations of relatively thin beds composed of mudstone,
sandstone and limestone are interpreted as cyclic changes
of depositional conditions (climatically driven?). Repeated
coarsening-upward successions with a transition from mudstone
to sandstone and/or finally limestone can be interpreted
as parasequences (Fig. 1).
In the Kralice area (Zágoršek et al. 2007, 2009; Fig. 1),
a carbonate production event is manifested by deposition
of calcareous mudstones to muddy limestones. The sediment
can be interpreted as having been deposited in a shallow
marine setting. The absence of sedimentary structures
induced by waves or tidal activity may point to offshore
conditions. Shells were eroded in coastal environments,
transported offshore by gravity currents or storm-induced
flows (tempestites), and rapidly deposited. Horizontal
lamination in mudstone points to the pulsed nature of sedimentation.
Intercalations of poorly sorted sand with angular
psefitic clasts, intraclasts of sandy clay or clayey sand
originating from the underlying bed, and shell debris with
abundant Rhodophyta reveal a typical textural inversion
that can be explained by highly disparate sediment deposition/transport
environments. Transport of outsized isolated
angular clasts can be explained by rock falls (Nemec 1990),
Fig. 2  Biostratigraphic correlation of the sections studied
	 Facies (2014) 61:419
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a relatively dramatic relief of the basin floor and redeposition
of large clasts into shallow marine environments by
density currents.
In the large limestone bodies (Kroužek and Podbrˇežice,
Fig. 1), three lithofacies were macroscopically identified. The
first is detrital limestone with reworked lithoclasts and bioclasts
(biodetrital grainstone). The amount of detrital material
varies both in volume and in grain size. The second lithofacies
reveals a very limited siliciclastic admixture; bioclasts
are separated by a sparitic or micritic groundmass (rudstonefloatstone).
Ripple cross-stratification and grading were the
most common sedimentary structures and were identified
especially within grainstones. The third lithofacies is calcareous
marl to sandstones. These “clastic intercalations” are only
thin interbeds within the dominant limestones. The lithology
and sedimentary structures indicate dynamic changes in depositional
conditions. Alternations of relatively calm periods
and predominant deposition from suspension with periods
characterised by either tractional or gravity currents can be
supposed.
Biofacies
The main allochems in the samples are bryozoan colonies,
coralline algae, unsorted allochems, lithoclasts and mollusks
with minor foraminifera and serpulids; the groundmass
consists of micrite and sparite (Fig. 3; Table 1). The
skeletal assemblage can be classified as heterozoan (James
1997) or as rhodalgal (Carannante et al. 1988).
Three microfacies were identified based on skeletal components,
the volume of lithoclasts and the mean grain size.
Compositions of each microfacies are summarized in Fig. 3
and Table 1. The microfacies are: (1) sandy coarse-grained
coralline algal–bryozoan–mollusc limestone (Fig. 4a); (2)
sandy bryozoan–coralline algal limestone (Fig. 4b); and (3)
bryozoan to bryozoan–coralline algal rudstone to floatstone
with grainstone matrix (Fig. 4c–d).
Differences in the fossil content of siliciclastic and mixed
siliciclastic‑carbonate units
Based on the occurrence of Helicosphaera waltrans, the
Mid Badenian sediments (sensu Hohenegger et al. 2014)
of the Carpathian Foredeep in Moravia can be divided into
two intervals (Švábenická 2002). In the older interval containing
H. waltrans, carbonates were not recorded; in the
Fig. 3  Composition of microfacies in the Podbrˇežice and Kroužek
sections. Ranges of component percentages in individual microfacies
are summarized. a Coarse-grained coralline algal–bryozoan–mollusc
sandy limestone; b Sandy bryozoan–coralline algal limestone; c Bryozoan
to bryozoan–coralline algal rudstone to floatstone with grainstone
matrix
Fig. 4  Microfacies (a–d) and coralline algae (e–h). a Coarsegrained
coralline algal–bryozoan–mollusc sandy limestone, Kroužek.
Limestone composed of protuberances and debris of coralline algae
(appearing as black) and elongated mollusk shells; b Sandy bryozoan–coralline
algal limestone, Kroužek. Sample consists mostly of
bryozoan debris with fewer coralline algae; c–d Bryozoan to bryozoans–coralline
algal rudstone to floatstone, Podbrˇežice. Bryozoan
colonies float in a grainstone to micritic matrix and are encrusted
with coralline algae; e Sporolithon lvovicum (Maslov) Bassi, Braga,
Zakrevskaya and Petrovna-Radionova; f Lithothamnion roveretoi Airoldi;
g Lithophyllum sp. 1; h Spongites fruticulosa Kützing
▸
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younger interval without H. waltrans, carbonates occurred.
To define the specific paleoceanographic situation during a
carbonate production event, firstly, the general differences
between the intervals with and without the deposition of
carbonates were tested using a Kruskal-Wallis test (Fig. 5).
The test showed that the units differ in relative abundances
of textulariids, epiphytic and oxyphilic benthic foraminifera,
Globigerina praebulloides and reworked calcareous
nannoplankton (higher in the siliciclastic-carbonate unit),
an abundance of calcareous nannoplankton and a relative
abundances of infauna, high-nutrient benthic foraminiferal
markers and Helicosphaera spp. (lower in the siliciclasticcarbonate
unit). Visual estimation (Fig. 5) showed higher
variances in abundances of Turborotalita quinqueloba, Par‑
agloborotalia mayeri, Reticulofenestra minuta and highnutrient
markers in the siliciclastic-carbonate unit.
Planktonic assemblages in the siliciclastic‑carbonate unit
In the siliciclastic-carbonate unit, planktonic organisms are
generally rare and only calcareous plankton was recorded.
However, siliceous plankton (diatoms and radiolaria) was
mentioned from isochronous siliciclastics of the Carpathian
Foredeep (Řeháková in Papp et al. 1978; Sláma 1983); in
the sections studied, they were recorded only below the
first limestone bodies in the Židlochovice boreholes.
Calcareous nannoplankton The Kruskal–Wallis test documented
statistically significant differences in nannoplankton
abundance between the limestones and siliciclastics in
the siliciclastic-carbonate complex (Fig. 6; Table 2): abundances
are approximately two times lower in the limestone
bodies than in the siliciclastics. In the former, peaks of calcareous
nannoplankton abundance can be correlated with
siliciclastic intercalations (Fig. 6).
Calcareous nannoplankton assemblages contain a total
of 15 Miocene species, of which Reticulofenestra minuta,
R. haqii, Coccolithus pelagicus and Thoracosphaera
spp. are common to abundant. The Kruskal–Wallis tests
showed statistically significant differences in the abundance
of Thoracosphera spp., which is higher in the limestone
bodies and of Reticulofenestra minuta which are
higher in siliciclastic intercalations within the limestones
(Fig. 6; Table 2). Though the assemblages look generally
very similar, dominated by Reticulofenestra minuta, nonmetric
multdimensional scaling (n-MMDS) showed the
differences between assemblages from a larger limestone
body (Kroužek and Podbrˇežice) and from an area dominated
by siliciclastics (Židlochovice and Kralice). Comparison
of multivariate statistical classification of samples
with relative abundances of the most common taxa
showed that variable but generally higher abundance of
Thoracosphaera spp. characterize assemblages from the
larger limestone body. Assemblages from the Židlochovice
and Kralice areas are less diverse with abundant Reticu‑
lofenestra minuta and variable abundance of Coccolithus
pelagicus. A higher abundance of Reticulofenestra haqii
is recorded in the Židlochovice area; Braarudosphaera
bigelowi appears in the Kralice area (Fig. 7).
Planktonic foraminifera (Fig. 8z, ab) The Kruskal–
Wallis test showed the differences between planktonic
foraminiferal assemblages in the limestone bodies, siliciclastics
and siliciclastic intercalations to be statistically
significant. These differences concern the P/B-ratio and the
abundance of cool-water/high nutrient plankton (Globige‑
rina spp. and Turborotalita quinqueloba), Globorotalia
spp. and Turborotalita quinqueloba (higher in siliciclastics,
lower in limestone bodies; Fig. 6). Of note, the P/Bratio
in limestones exceeds 10 % mainly in the siliciclastic
intercalations. nMMDS as well as relative abundances
of the most common taxa showed variable compositions
of assemblages. Assemblages from the Podbrˇežice section
substantially differ by a high abundance of Globigeri‑
noides spp. and Orbulina suturalis and a low abundance of
5-chambered Turborotalita quinqueloba and Globigerina
ottnangiensis. Assemblages from the Židlochovice and
Kralice boreholes are characterized by a higher abundance
of globorotaliids; a scarcity of four-chambered Globigerina
spp. was recorded in the Židlochovice area (Fig. 9).
Benthic biota in the siliciclastic‑carbonate units
Coralline algae (Fig. 4e–h) Lithothamnion roveretoi Airoldi,
Lithothamnion sp. 1, Mesophyllum cf. printzianum Woelkerling
& Harvey Adey, Lithophyllum sp. 1, and Spongites
fruticulosa Kützing were identified in the Podbrˇežice section
(Hrabovský et al. submitted). Sporolithon lvovicum
(Maslov) Bassi, Braga, Zakrevskaya & Petrovna-Radionova,
Lithothamnion sp. 2, Mesophyllum curtum Lemoine, M.
cf. printzianum Woelkerling & Harvey, Phymatolithon sp.,
Lithophyllum sp. 1, L. sp. 2, Hydrolithon lemoinei (Miranda)
Aguirre, Braga & Bassi, and Spongites fruticulosa Kützing
were identified in the Kroužek section (Hrabovský et al. submitted).
According to these authors Mesophyllum cf. print‑
zianum and Spongites fruticulosa inhabit present-day inner
shelf environments with normal salinity.
A diverse assemblage comparable to that at Kroužek was
documented in Židlochovice (Hrabovský et al. submitted).
Benthic foraminifera (Fig. 8a–y, ac–ad). The abundances
of benthic foraminifera in limestones vary from 50 to 700
Fig. 5  Differences in the relative abundance of groups of microfossils
between the siliciclastic and siliciclastic-carbonate units. Statistically
significant differences (tested by Kruskal–Wallis test) are
marked by light yellow arrows (p value from 0.05–0.0001) or dark
yellow arrows (p value <0.0001). SC siliciclastic unit with H. wal‑
trans; S-CC siliciclastic-carbonate unit above the LO of H. waltrans;
p- p-values for tested parameters
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specimens/g of rock; values of 700–1,000 specimens/g
were reached in the siliciclastic intercalations in the
Podbrˇežice section. In the siliciclastics, there abundances
may reach more than 1,000 specimens/g. Kruskal–Wallis
tests showed statistically significant differences in the
composition of benthic foraminiferal assemblages (Fig. 6;
Table  2): siliciclastics differ from limestones in higher
abundances of infauna, high-nutrient markers and textulariids,
in the lower values of BFOI (=the Benthic Foraminiferal
Oxygenation Index, Kaiho 1994), and in lower abundances
of epiphytic foraminifers and Elphidium spp.
Siliciclastic intercalations in limestones are characterized
by higher abundances of Ammonia spp.
Generally, the high abundances of Asterigerinata
planorbis (40–60 %) are very characteristic for all samples
from the limestones bodies. In limestones, three groups of
benthic foraminifera species with statistically significant
differences of relative abundances based on Spearman
correlation coefficients can be distinguished (Fig. 10): (1)
Bulimina spp., Bolivina spp., Hansenisca soldanii, Melo‑
nis pompiloides, Cassidulina spp., Globocassidulina spp.,
lagenids and textulariids; (2) Ammonia spp., Elphidium
spp., and Nonion commune; (3) Cibicidoides spp. and
Lobatula lobatula. The nMMDS (Fig. 11) clearly separated
assemblages from the Kralice sections due to the higher
abundances of taxa from group (1), mainly Cassidulina
spp. Assemblages from the limestones of the Židlochovice
section can be characterized by dominance of taxa from
groups (2) and (3), Asterigerinata planorbis (common)
and at some levels higher abundances of Bolivina dilatata,
Globocassidulina spp., and Cassidulina spp. from group
(1). Assemblages from the Podbrˇežice and Kroužek sections
are very similar and differ from the assemblages of
Fig. 6  Differences in quantitative parameters of microfossil assemblages
among siliciclastics (CLT), carbonates (LST) and siliciclastic
intercalations in limestones (ICL) in the siliciclastic-carbonate unit.
Statistically significant differences (tested by Kruskal–Wallis test) are
marked by yellow arrows. P values are summarized in Table 2. BFOI
the benthic foraminiferal oxygenation index; ratio between oxyphilic
and hypoxic benthic foraminifera
Table 2  Kruskal–Wallis test
p values: evaluated differences
in quantitative parameters of
microfossil assemblages among
siliciclastics, carbonates and
siliciclastic intercalations in
limestones of the siliciclasticcarbonate
unit
Characteristics Limestones/
intercalations
Limestones/
elastics
Clastics/
intercalations
Benthic forminifera
 Foraminiferal number 0.00945 0.01822 0.5916
 Agglutinated 0.3894 0.0037 0.000704
 BFOI 0.9545 2.8E−9 2.29E−7
 Infauna 0.785 2.63E−8 1.06E−6
 High-nutrient 0.765 1.02E−8 4.41 E−8
 Epiphytic 0.2009 3.66E−9 4.03E−6
 Euryhaline 0.008256 0.00119 5.14E−5
 Elphidium spp. 0.1666 2.78E−6 1.40E−5
 Ammonia spp. 0.2015 0.0635 0.01594
 Large 0.2455 0.08701 0.2553
Planktonic forminifera
 P/B-ratio 0.403 1.33E−8 7.39E−5
 Turborotalita quinqueloba 0.2875 5.96E−7 1.73E−8
 Small Globigerina spp. 0.8907 4.05E−8 0.00015
 Globigerina praebulloides 0.1306 1.65E−6 3.41 E−5
 Cool-water/high-nutrient 0.8137 7.48E−7 0.000139
 Warm-water/low-nutrient 0.7173 0.001562 0.000501
 Globigerinoides spp. 0.3765 0.197 0.8912
 Orbulina spp. 0.9011 0.142 0.2554
 Globorotalia spp. 0.6837 2.96E−6 4.37E−7
Calcareous nannoplankton
 Calcareous nannoplankton abundance 0.01537 1.26E−6 5.31 E−5
 Coccolithus pelagicus 0.5828 0.1381 0.5193
 Reticulofenestra minuta 0.4382 0.1391 0.04827
 Reticulofenestra haqii 0.229 0.9195 0.3475
 Thoracosphaera spp. 0.03226 1.51E−9 4.13E−6
 Cyclicargolithus floridanus 0.2261 0.8105 0.2788
◂
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the Židlochovice and Kralice areas by abundant Asterigeri‑
nata planorbis and the absence of taxa from group (1). The
assemblages of Kroužek section and of the middle part of
Podbrˇežice section differ from the other ones at Podbrˇežice
only in slightly higher abundances of small-sized cibicidoids
and Lobatula lobatula.
Molluscs The molluscan association at Kroužek is relatively
rich; it includes species of Corbula, Ostrea, Turri‑
tella, Conus, Nassarius, Petaloconchus, Lemintina, Gen‑
ota, Bittium, etc. At Židlochovice, the molluscan fauna
consists predominantly of bivalves; gastropods are less
common. Small gastropods of the genera Bittium, Alvania,
Solariorbis, Gibbula, and Rissoina were identified in some
Fig. 7  a–d Differences in the calcareous nannoplankton assemblages using non-metric multdimensional scaling (nMMDS); e–h Comparison of
relative abundances of common taxa; i Location of sections studied in basin; j Shepard plot of nMMDS
Fig. 8  Foraminifera. a–e Asterigerinata planorbis (d’Orbigny),
Kr 21, Pr 21, Kr 21, Pr 9, Kr 9; f–l Small-sized Cibicidoides spp.,
Kr 21 Kr 17, Kr 7, Kr 12, Kr 12, Kr 9, Kr 22; m Lobatula lobatula
Walker and Jacob, Kr 23; n Cibicidoides austriacus (d’Orbigny),
Kr 23; o Quinqueloculina akneriana d’Orbigny, Kr 23; p Elphidium
flexuosum (d’Orbigny), Kr10; q Elphidium subtypicum Papp, Kr 22;
r Elphidium fichtelianum (d’Orbigny), Kr 2; s Elphidium macel‑
lum (d’Orbigny), Kr 7; t Nonion sp., Pr 21 undeterminable corroded
and broken test; u Uvigerina acuminata Hosius, Pr 14; v Bolivina
plicatella Cushmann, abraded test, Pr 10; w Pullenia bulloides
(d’Orbigny), Pr 14; x Melonis pompiloides (Fichtel and Moll), Kr 17;
y Ammonia viennensis (d’Orbigny), abraded test, Pr 20; z–aa Glo‑
bigerinoides bisphericus Todd, 26-Kr 7, 27, Kr 22; ab Globorotalia
peripheroronda Blow and Banner, Kr22; ac Pararotalia aculeata
(d’Orbigny), Kr 10; ad Amphistegina bohdanowiczi Bieda, Kr 10.
Lenght of scale bars 100 µm
▸
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	 Facies (2014) 61:419
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intervals. At Kralice, the mollusc fauna is composed predominantly
of bivalves; fragments of pectinids dominate,
chitons are rare, and gastropods are completely absent.
Palynomorphs
From a palynological point of view, all samples from
limestone bodies (with sand intercalations) were sterile.
These sediments were not suitable for the preservation of
palynomorphs due to their large grain size (without clay
admixture) and chemical conditions during sedimentation.
Pollen grains were probably washed out from these
sediments and destroyed by oxidation. The absence of
pollen grains in ancient carbonate environments was also
described by Heusser (1978).
The palynomorph content seemed to be also dependent
on the redox potential in the siliciclastics (Heusser
1978; Doláková et al. 2014). Large segments of the ageequivalent
sediments from the boreholes in Židlochovice,
Rebešovice, Chrlice, Opatovice and Oslavany were
Fig. 9  a–d Differences in the planktonic foraminifera assemblages using non-metric multdimensional scaling (nMMDS); e–h Comparison of
relative abundances of common taxa; i Locations of sections studied in the basin; j Shepard plot of nMMDS
Facies (2014) 61:419	
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studied (Hladilová et al. 1999; Doláková et al. 2011;
Doláková et al. 2014). They revealed periodic changes of
oryctocoenoses with diversified pollen spectra followed by
a strong dominance of conifers together with marine dinoflagellates
and, afterward, the disappearance of all pollen
and spores. Above that succession, limestone layers were
recorded.
Flora with up to 30 % of evergreen, broad-leaved forest
elements (Engelhardia, Platycarya, evergreen Fagaceae:
Castanopsis, Trigonobalanopsis) is thought to have a
mainly subtropical character. The share of warm-temperate
mixed-mesophytic and broad-leaved deciduous forest
members (i.e., Quercus, Carya, Celtis, Juglans) was
lower. A higher diversity of “oak type” pollen grains (e.g.,
Quercus robur-pubecscens) and the general occurrence
of Platanus pollen were typical. The proportions of most
thermophilous elements of P1 sensu Mai (1981, 1991) and
Stuchlik (1994) slightly decreased in comparison with the
Lower Miocene palynospectra of the Carpathian Foredeep
(Doláková et al. 2011; Kovácˇová et al. 2011). The abundant
and diverse Pinaceae (with an admixture of Cedrus, Abies,
Picea, Cathaya and Tsuga) were components of mountain
conifer-rich forests.
Fluctuation of coastal swamp and riparian elements
could be a result of humidity changes. Herbs and heliophilous
elements such as Poaceae, Asteraceae, Caryophyllaceae,
Chenopodiaceae, Olea, Buxus and Ephedra indicate
the existence of more open areas and floral elements growing
on drier places at this time. Warm-wet conditions, an
increase in seasonality and cooler phases were observed
within the subtropical character of the terrestrial flora
(Doláková et al. 2014).
Oxygen and carbon isotopes
Stable oxygen and carbon isotopic values from bulk samples
of limestones and siliciclastic intercalations from a large
limestone body (“Rousínov ridge”) were compared (data
from Nehyba et al. 2008b). The Kruskal-Wallis test showed
a statistically significant difference between δ18
O and δ13
C
values in the siliciclastic intercalations of the Kroužek section
(Fig. 12a). Kroužek limestones are more δ13
C-enriched
Fig. 10  Statistically significant correlations (p values < 0.05) among
relative abundances of benthic foraminifera in limestones: three
groups of species are distinguished by different colors of circles. Positive
values of Spearman coefficient are given in wide arrows, negative
values next to slender arrows
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than those from Podbrˇežice. In the Židlochovice boreholes
(Fig. 12b), the planktonic foraminifers have lower δ13
C and
δ18
O values relative to their benthic counterparts. This difference
in δ13
C values is enhanced for the limestone-hosted
tests relative to those collected from siliciclastic layers. At
some levels (ZIDL1/3.1 m; ZIDL2/12.9 m), the isotopic
differences between planktonic and benthic foraminifera
are minimal. Contrary to the Židlochovice data, in the
siliciclastics from the Kralice section (Fig. 12c), carbon isotope
compositions of foraminifers are significantly lower
in planktonic tests than in benthic ones, while the isotopic
oxygen difference between plankton and benthos is only
slightly lower. In these limestones, the isotopic carbon compositions
of these foraminifer types overlap, while δ18
O values
are higher for plankton, opposite to samples from the
Židlochovice boreholes.
Fig. 11  a–d Differences in benthic foraminifera assemblages using non-metric multidimensional scaling (nMMDS). e–h Comparison of relative
abundances of common taxa. i Location of studied sections in basin. j Shepard plot of nMMDS
Fig. 12  a Differences in isotopic oxygen and carbon values of bulk
sediment between the Kroužek and Podbrˇežice sections; b Isotopic
oxygen and carbon values of foraminifera tests in the ZIDL-1 borehole;
c Isotopic oxygen and carbon values of foraminifera tests in the
Kralice section
▸
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	 Facies (2014) 61:419
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Discussion
Correlation of the Middle Miocene carbonate production
event across the Central Paratethys
A carbonate production event, manifested by the deposition
of a mixed carbonate–siliciclastic unit in the Moravian part
of the Carpathian Foredeep is dated by the co-occurrence
of Praeorbulina, Orbulina and Sphenolithus heteromor‑
phus, and the absence of H. waltrans. A similar biostratigraphic
position of the siliciclastic-carbonate unit was also
recorded in the Ugljevic section (Pezelj et al. 2013). In the
Transylvanian Basin, the studied limestones contain only
Orbulina sp. (Filipescu and Gîrbacea 1997; Zágoršek et al.
2010); nevertheless, nannoplankton assemblages enable
correlation with the upper part of the NN5 Zone without H.
waltrans (Holcová, unpublished data). Croatian limestones
from the Zrin Basin (Martinuš et al. 2012) can be correlated
with the NN5 Zone; H. waltrans was not recorded. The cooccurrence
of Praeorbulina and Orbulina is characteristic
for the appearance of limestones in the Mailberg Formation
of the Eastern Alpine Foredeep in Austria (Mandic
2004). The formation correlates with chron C5Bn.r, based
on paleomagnetic data from C´orić et al. (2004). The age of
the limestone bodies in Hungary, Poland and Ukraine (Pisera
and Studencki 1989; Randazzo et al. 1999; Radwan´ski
et al. 2006) cannot be determined so strictly; nevertheless,
the same age is very probable. However, other younger
intervals involving carbonate production in the Badenian
were described from Ukraine (Gorka et al. 2012).
Despite these uncertainties, the carbonate production
event in the Central Paratethys can be correlated with the
NN5 Zone. While magnetostratigraphic dating in the Eastern
Alpine Foredeep (chron C5Bn.r, based on paleomagnetic
data; C´orić et al. 2004) enumerated the age at 15.034–
14.888 Ma, the biostratigraphic events dated the carbonate
production in the Moravian part of the Carpathian Foredeep,
in the Transylvanian Basin, and in the Zrin Basin to
the upper part of the NN5 Zone, above the LO of H. wal‑
trans at approximately 14.4–14.3 Ma (the LO of H. wal‑
trans) to 13.42 Ma (the LO of S. heteromorphus).
Factors triggering the Middle Miocene carbonate
production event
The start of the mixed siliciclastic-carbonate sedimentation
was connected with the following paleoenvironmental
changes: (1) Decrease of siliciclastic input in the basin;
the decrease can be indirectly interpreted from the disappearance
of siliceous plankton, which may be caused by
the lack of a silica source. The silica in the Early-Middle
Badenian deposits of the Carpathian Foredeep could originate
from distal Badenian acidic air fall tephra or from
the weathered crystalline basement along the passive,
i.e., western, margin of the basin. The strongly restricted
occurrence of volcaniclastic beds in the successions studied
points to the dominant role of a decreased input of
reworked and weathered siliciclastics.
The reduction of siliciclastic input can be connected
either to transgressive and early highstand conditions, aridification
of climate, or to both. The beginning of the Middle
Miocene was characterized in the Carpathian Foredeep by a
large marine transgression affecting the entire circum-Mediterranean
area (Rögl 1999; Popov et al. 2004; Kovácˇ et al.
2007; Piller et al. 2007). The carbonate production event is
connected to the culmination of this transgression, which is
documented by the occurrence of the denudation relicts of
siliciclastic-carbonate deposits far on the Bohemian Massif
(Hladilová and Zdražílková 1989). Thus, deposition of the
carbonate–siliciclastic complex corresponds to the maximum
flooding zone.
The Langhian aridification events were described from
the Mediterranean area (Hüsing et al. 2010). The first discrete
paleoenvironmental step, at 15.074 Ma, might be
linked to reduced riverine discharge and climatic cooling
(Hüsing et al. 2010). The dating of this step agrees with
the appearance of carbonate bodies in the Alpine Foredeep
(Mandic 2004). The second discrete step occurred at
14.489 Ma. This age corresponds to the onset of carbonate
sedimentation in the Carpathian Foredeep. Findings
of pollen grains of herbs and heliophilous plants such as
Poaceae, Asteraceae, Caryophyllaceae, Chenopodiaceae,
Olea, Buxus and Ephedra indicate the existence of more
open and dryer areas at that time; (2) The appearance of
sea-grass meadows and a well-aerated sea-floor can be
interpreted from the statistically significant increase in the
abundance of epiphytic as well as oxyphilic taxa. Pyrite
infillings of foraminiferal tests, described from the older
siliciclastic interval (Tomanová-Petrová and Švábenická
2007), disappeared, which also indicates an increase of
oxygenation. This may be correlated with the establishment
of downwelling (anti-estuarine) circulation (Brzobohatý
1987; Báldi 2006) and the culmination of the third order
transgression, connected with the appearance of a shallow
marginal part of the basin suitable for the expansion
of sea-grass meadows. Seagrasses are, indirectly, important
producers of biogenic CaCO3 because their epibionts,
invertebrate shells and coralline algae occur often in high
density (Brasier 1975). Sea-grass respiration and photosynthesis
cause variations in the O2 and CO2 content of seawater,
which influences the rate of CaCO3 fixation by marine
organisms (Davies 1970). (3) Environmental instability
and an increase in seasonality can be interpreted from the
variable abundances of the opportunistic Reticulofenestra
minuta. It indicates environmental stress in the upper part of
the water column, characterized by fast changes within that
Facies (2014) 61:419	
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environment, including fluctuations of salinity from brackish
to hypersaline (Wade and Brown 2006), and oscillations
of the nutrient content between oligotrophic and eutrophic
(Wells and Okada 1997; Flores et al. 1997; Kameo 2002).
Instability can also be interpreted from the large variation
in the abundance of Turborotalita quinqueloba, a marker of
cold, non-stratified water masses (Hohenegger et al. 2008),
and Globorotalia mayeri. The large abundance variability
of high-nutrient markers in the benthic assemblages shows
the instability of environmental conditions at the sea floor.
The bimodal abundance distributions of all these groups
may indicate short-term (maybe seasonal) changes of nonstratified,
mixed eutrophic waters with small Turborotalita
quinqueloba and high nutrient markers at the bottom; the
stratified water column is marked by oligotrophic warmer
water with Globorotalia mayeri in the upper part. Salinity
fluctuations in the upper part of water column cannot
be excluded (Kendl, pers. comm., determined dinoflagellate
markers of hypersaline conditions). The alternation of
stratified and mixed water columns is also confirmed by the
oxygen and carbon isotope data (Fig. 12).
The instability (probably connected with increased seasonality)
can be connected with environmental and climatic
changes preceding the Middle Miocene Climate Transition
(MMCT) at 13.82 Ma (Holbourn et al. 2005). In terrestrial
climates, the MMCO/MMCT transition is characterized by
an increase in the mean annual range of temperature, mainly
due to decreasing cold month temperatures (Bruch et al.
2010); increased seasonality is principally expressed in the
seasonality of precipitation (Doláková et al. 2014). While
the decrease of terrigenous input and oxic conditions with
seagrass meadows accelerated the production of carbonates,
the instability of the environment was a rather unfavorable
factor. It could have caused the reduced thickness of limestone
bodies and high amount of siliciclastic components
in the carbonates, indicating only short-term and spatiallyrestricted
intervals suitable for carbonate deposition.
Cyclicity in the mixed limestone‑siliciclastic complex
Orbital forcing climatic cyclicity strongly influenced the
evolution of the Middle Miocene ecosystems in the Central
Paratethys (Hohenegger et al. 2009) and affected the paleobiological,
as well as the geochemical and sedimentological,
characteristics of sediments. Cyclical changes in the
mixed siliciclastic-carbonate units are manifested by the
alternation of paleoenvironments suitable for the deposition
of limestones and siliciclastics. The direct relation between
orbital forcing climate changes (wet-dry climatic cycles)
and pelagic limestone-marl alternation was documented in
the Langhian succession of the Conero Riviera by Mader
et al. (2004). According to these authors, limestones represent
dry/colder-water periods and marls represent wet/
warmer-water periods with limited productivity and
increased terrigenous supply.
To test the character of cyclical changes in the study area
and compare the local model with the Mediterranean one,
the paleoenvironments represented by the limestones and
siliciclastics in the mixed limestone-siliciclastic unit were
interpreted separately.
Paleoenvironment during deposition of limestones
Generally, the limestones in the Central Paratethys are
interpreted as warm-water (Filipescu and Gîrbacea 1997;
Pouyet and Tarkowski 1998; Riegl and Piller 2000;
Radwan´ski et al. 2006; Basso et al. 2008; Doláková et al.
2008; Martinuš et al. 2012; Pezelj et al. 2013). However,
Randazzo et al. (1999) considered the limestones from the
Pannonian Basin as being cool-water. This interpretation
does not agree with other paleoclimatological data from
the Central Paratethys (Kvacˇek et al. 2006; Doláková et al.
2014). The estimated paleotemperatures agree with a subtropical
climate, which is expected from the biofacies analysis
of red algal limestones (Kroeger et al. 2006).
In agreement with results from the other Central Paratethys
basins, rhodalgal or bryorhodalgal facies dominate
within the Early‒Middle Badenian limestones (Pisera and
Studencki 1989; Randazzo et al. 1999; Studencki 1999;
Radwan´ski et al. 2006; Doláková et al. 2008; Martinuš
et al. 2012; Wiedl et al. 2012). Using recent analogues from
the Mediterranean (Martinuš et al. 2012), subtropical, lowto
moderate-energy clean waters with a depth of 10–80 m
(mainly 20–40 m) are expected during deposition of these
types of carbonates.
Mollusc assemblages from Kroužek correspond well
to this microfacies interpretation. A simple paleoecological
analysis (Hladilová 1984) generally indicated a shallow
(littoral to sublittoral, with a maximum depth of up to
80 m) warm-water environment of marine salinity (above
30 ‰), good aeration, sufficient light and a prevailing soft
bottom. Variations in water energy can be observed.
Recorded mollusc and foraminifera species are suspension-feeders,
detritivores, predators and herbivores. The
ratio between these trophic groups in assemblages varies
but suspension-feeders, herbivores or predators always predominate
over detritivores and deposit-feeders. This indicates
a deficiency of organic detritus in the sediment as a
nutrient source during deposition of the limestones.
During the deposition of limestones in the Podbrˇežice
area the upper water column was characterized by a low
abundance of plankton. The presence of Globigerinoides
spp. indicates oligotrophic and well-stratified waters (Reynolds
and Thunell 1985; Hemleben et al. 1989), and the characteristic
nannoplankton genus, Thoracosphaera spp., is
a general proxy for oligotrophy or stratification (Höll et al.
	 Facies (2014) 61:419
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419  Page 20 of 26
1998; Vink et al. 2002). Thus, a stratified water column with
oligotrophic conditions in the upper layer can be expected
during the deposition of limestones, also confirmed by the
absence of Turborotalita quinqueloba. The latter occurs in
siliciclastics and is a marker of a cold, non-stratified water
body. However, autochthony of the planktonic foraminifera
in the limestone bodies is questionable. Transport of plankton
is confirmed by the assumed depth for Asterigerinata
assemblages (up to 100 m; Holcová and Zágoršek 2008), in
which live planktonic foraminifera are extremely rare. The
transport mechanism of planktonic foraminifera from central
to marginal parts of the basin may have been expected
downwelling circulation (Brzobohatý 1987; Báldi 2006).
The discrepancy between downwelling circulation leading
to mixed water masses and presence of markers of a stratified
water column may be explained by seasonal changes of
the circulation pattern, with downwelling in summer and a
stratified water body in winter, which is in agreement with
the data of Hohenegger et al. (1999) and others. Seasonality
is supported by the variable oxygen isotope composition of
benthic foraminifera which may indicate mixing of seasonal
populations (Holcová and Demeny 2012).
The increase in oxygen values in the Kralice section at
the transition from siliciclastics to limestones and the oxygen
isotope composition of bulk sediment from the limestone
bodies in the “Rousínov Ridge”, stratigraphically
higher than that of isochronous data from the Mediterranean
(Mader et al. 2004; Fig. 12a), may indicate higher
evaporation in the marginal part of the Central Paratethys
basin and a decrease of temperature. Higher evaporation is
supported by assemblages composed only of miliolids in
the upper part of the Kroužek section. The absence of coral
patch reefs in the Moravian part of the Carpathian Foredeep,
which are known from isochronous rocks in Austria,
Hungary, Ukraine and Poland (Pisera and Studencki 1989;
Randazzo et al. 1999; Radwan´ski et al. 2006; Martinuš
et al. 2012; Wiedl et al. 2012), may be also due to both
fluctuations in salinity and lower temperatures. Salinity
fluctuations are more likely because the Polish and partly
Ukrainian basins were located at a higher latitude than
Moravia (Popov et al. 2004). Although it is not possible to
exclude the influence of lower temperatures on the oxygen
isotope values and on the absence of coral patch reefs during
deposition of limestones in the Carpathian Foredeep,
increasing salinity appears to have been the decisive factor.
Paleoenvironment during deposition of siliciclastics in the
limestone‑siliciclastic unit
During deposition of siliciclastics, both planktonic and
benthic assemblages showed enhanced productivity. In the
upper layer of the water column, the abundance of calcareous
nannoplankton and planktonic foraminifera increased.
The increase of small, 5-chambered Turborotalita quinque‑
loba indicates a non-stratified water column characterized
by intensive vertical mixing or upwelling (Reynolds and
Thunell 1985; Hemleben et al. 1989). Globorotaliids, as is
the case with other abundant groups of planktonic foraminifera,
are classified as cold-temperate taxa (Bicchi et al. 2003)
and indicate surface water stratification (Pérez-Folgado et al.
2003). Alternation of assemblages dominated by T. quinque‑
loba and by Globorotalia indicates that the alternation of
stratified and mixed water bodies persisted from the period
of carbonate production. The increase in nutrient content at
the bottom, indicated by a higher abundance of high-nutrient
markers and detritivores (textulariids and deeper infauna), is
connected with the appearance of organic detritus in the sediment.
The lower Benthic Foraminiferal Oxygenation Index,
as well as the disappearance of oxyphilic epiphytic taxa,
showed a decrease of oxygen content at the bottom.
There are two possible ways to explain the nutrient sources:
terrigenous input or seasonal upwelling. The decrease in carbon
isotope values of planktonic foraminifers from the siliciclastics
in the marginal area of the basin (Kralice section) in
comparison with the values from the central part of the Carpathian
Foredeep (which are near to the values from the Mediterranean
Sea and Atlantic Ocean) favours terrestrial input.
Furthermore, the occurrence of Cassidulina as a marker of
phytodetritus supply and its decrease from marginal (Kralice)
to more central parts of the basin suggests input of nutrients
from the continent. Thus, deposition of siliciclastics corresponded
with a more humid climate with higher rainfall.
In the Zrin Basin (Pezelj et al. 2013), high-nutrient markers
(Bulimina, Globocassidulina and Valvulineria) occur
also in siliciclastics. The alternation of siliciclastic and carbonate
sediments can be explained by sea-level fluctuations
(Mandic 2004) in the Eastern Carpathian Foredeep: the marly
layers are characterized by an enhanced richness and heterogeneity
of benthic foraminifera and by abundant planktonic
foraminifera. These indicate open-marine conditions of the
upper middle shelf, while carbonate production might be
linked with shallowing. To summarize, the genesis of the
mixed limestone-siliciclastics unit can be explained by alternating
wet and dry phases: during the wet phase, siliciclastics
were deposited due to higher terrigenous input. Carbonates
were deposited during the dry intervals, connected with
low nutrient input and at least occasional increases in salinity.
For both intervals, short-term (probably seasonal) changes
between mixed and stratified water masses are assumed.
Spatial paleoenvironmental variations during times
of carbonate deposition
Though generally limestones were deposited in a uniform
environment, some differences among individual areas
existed:
Facies (2014) 61:419	
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1.	 In the Kralice Bay (Zágoršek et al. 2009; Holcová and
Demeny 2012), sedimentation of carbonates is connected
with the regressive phase of fourth or lower
orders of sea-level changes. Carbonates contain a
high admixture of siliciclastics and indicate rather
unfavourable conditions for carbonate production.
A facies study shows the pulsed nature of sedimentation,
a relatively dramatic relief of the basin floor and
redeposition of clasts to shallow marine environments
by density currents. Molluscs represent a mixture of
different environments, which also indicates higher
water energy. Bivalves, as well as benthic foraminiferal
assemblages, differ from other sections by higher abundances
of detritivores and suspension feeders, confirming
the environment to be rich in organic detritus and
suspended organic particles. Holcová and Demeny
(2012) interpreted the bay to be influenced by seasonal
phytodetritus supply connected with the bloom of
opportunistic taxa. However, the almost total absence
of brackish and estuarine elements, and the presence
of occasional brachiopods, confirm that the terrigenous
input did not decrease salinity, because brackish-water
biota have not been recorded either by autochthonous
or allochthonous taxa. The situation in the upper part
of the water column is very well comparable with
that of the “Rousínov Ridge”: planktonic foraminifera
are rare and very likely have been transported by
superficial currents from the central part of the basin.
More abundant and probably autochthonous planktonic
foraminifera were recorded in sample KRA8
with its higher oxygen isotope values; this may indicate
a decrease of temperature and/or increase in salinity.
Since a lower temperature in the upper part of the
water column is less likely, a higher rate of evaporation
in the marginal part of the basin is preferred. Higher
variations in benthic isotope values may have been
caused be greater seasonal differences. The occurrence
of Thoracospharea spp. indicates oligotrophic waters
and, together with different oxygen isotope values for
plankton and benthos, also indicates at least a seasonal
termohaline stratification of the water column.
2.	 Limestones from the Židlochovice site were deposited,
as at Kralice, during a sea-level lowstand (of
fourth or lower order; Doláková et al. 2014). Similar to
Kralice, the mixing of elements of the deeper infralittoral
and shallow sublittoral, or on rare occasions, even
with elements of an exposed rocky coastline, indicates
relatively high water energy; the almost total absence
of brackish and estuarine elements confirms that even
at the eastern coast of the Carpathian Foredeep brackish
biota did not exist or were rare.
	 Foraminiferal assemblages from Židlochovice differ in
their higher abundance of detritivores: infaunal Boliv‑
ina dilatata and markers of terrigenous input, Globo‑
cassidulina spp. and Cassidulina spp. The appearance
of Turborotalita quinqueloba indicates non-stratified
and the most eutrophic waters in comparison with
other areas of limestone deposition studied.
	 Variable isotope values may be explained by the allochthony
of some limestones. The isotope values from thin
limestone bodies correspond to values from siliciclastic
intercalations in the “Rousínov Ridge”; therefore, an
allochtonous origin of material is expected. Deposition
of allochthonous limestones may be connected with
comparable catastrophic climatic events, as assumed
for deposition of the siliciclastic intercalations.
3.	 The “Rousínov Ridge” limestones were deposited
during the whole fourth or lower order transgressiveregressive
cycle (Zágoršek and Holcová 2005). Various
environments at the “Rousínov Ridge” can be interpreted
mainly from microfacial differences that show
the decisive role of hydrodynamic conditions in the differentiation
of the limestone bodies. The Kroužek section
is characterized by more lithoclasts, molluscs, a
diverse assemblage of coralline algae and sparite, suggesting
a more dynamic and shallower paleoenvironment
with a higher input of lithoclasts in comparison
with Podbrˇežice, where a deeper and sheltered environment
existed. Also, a higher abundance of the opportunist
Cibicidoides sp. and of euryhaline foraminiferal
species indicate more stress and an unstable near-shore
environment with salinity fluctuations in the Kroužek
section in comparison with Podbrˇežice. Molluscs from
Kroužek differ from other sections in that bivalves are
dominant and gastropods occur only sporadically. This
dissimilarity can be explained both by primary differences
in the molluscan assemblages, and probably also
by a certain degree of sorting of shells. The primary differences
are probably related to water energy, depth and
substrate types. Specific conditions of the Podbrˇežice
sections indicate a facies with pedunculate bryozoans
and a high amount of micrite, recorded for the first time
from the Central Paratethys. Hageman et al. (2000)
interpreted the paleoecological differences between
rhodolith and rhodolith–bryozoan accumulations and
assumed that bryozoans and a higher abundance of micrite
indicate a deeper and colder environment, which is
supported by microfacies and foraminiferal data. The
statistically significant (Kruskal–Wallis test) increase
in the carbon isotope composition in Kroužek may also
have been caused by higher evaporation.
To summarize the individual interpretations, the
paleobiological, microfacies and sedimentological differences
in the individual carbonate bodies reflect variations
in the intensity of terrigenous input (higher in
	 Facies (2014) 61:419
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419  Page 22 of 26
Kralice and Židlochovice; lower on the “Rousínov
Ridge”), the environmental dynamics, depth (above
and below wave base) and morphology of the sea floor,
which was predisposed to gravity flows. These flows are
thought to occur mainly in the Židlochovice area, where
the majority of limestones most likely consist of allochthonous
components.
Genesis of siliciclastic intercalations within the limestones
Thin (cm-thick) siliciclastic intercalations were recorded
in the limestones bodies of the “Rousínov Ridge.” They
occur at irregular distances, from tens of centimetres to
two metres. The temporal frequency of these intercalations
is hard to estimate due to the unknown depositional rate.
Fig. 13  Paleoenvironmental model of the carbonate production event
Facies (2014) 61:419	
1 3
Page 23 of 26  419
Generally, the sedimentation rate of comparable types of
carbonates is high, but diagenetic compaction may substantially
decrease the thickness. However, an occurrence every
few hundred years is likely.
A high abundance of Ammonia sp. in these intercalations
(Fig. 6) accompanied by a low abundance of infauna
suggest a decrease in salinity during their deposition. A
higher abundance of Reticulofenestra minuta and higher
BFOI indicate stress conditions. In Podbrˇežice, an increase
in the abundance of the suspension-feeder Lobatula lobat‑
ula (Murray 2006) and detritivorous infaunal species, as
well as of plankton, both primary producers (nannoplakton)
and consumers (planktonic foraminifera) and, particularly
of small-sized Globigerina spp. (Zágoršek and Holcová
2005), indicate a high level of nutrients in suspension. In
the Kroužek section situated nearer to the coastline, siliciclastic
intercalations are more numerous and of a greater
thickness. Both oxygen and carbon isotope values are statistically
significantly lower in the intercalations in this
section (Fig. 12a). All these observations could indicate
that siliciclastic intercalations were deposited during high
episodic input of terrigenous material with nutrients connected
to the input of freshwater. This may correspond
with “extreme climate events” deduced from the strongly
negative excursions in δ18
O and δ13
C in the Ostrea isotope
archive (Harzhauser et al. 2011). The expected frequency
of siliciclastic intercalations of about several hundreds of
years corresponds with the assumed frequency of catastrophic
storm and rainfall events. As mentioned above,
deposition of the thin allochthonous limestone bodies at
the western coastline near Židlochovice may be related to
comparable catastrophic climatic events as the deposition
of siliciclastic intercalations.
Comparable intercalations have been described in the
Eastern Alpian Foredeep (Mandic 2004). Similar to our
interpretations, the disappearance of plankton, the decrease
of benthic foraminifera diversity and the dominance of
opportunistic Cibicidoides and euryhaline Elphidium in
these intercalations may have been caused by a salinity
decrease due to freshwater input.
Based on the paleoenvironmental interpretation presented
above, a paleoenvironmental model of a carbonate
production event has been compiled in Fig. 13.
Conclusions
1.	 A carbonate production event in the Central Paratethys,
connected with deposition of a siliciclastic-carbonate
unit in the marginal part of the basin, can be correlated
with the culmination of transgression in the NN5 Zone.
The event was caused by a decrease of terrigenous input
due to the high-stand conditions and, above all, aridification.
Aridification preceded the Middle Miocene Climate
Transition. The event can also be correlated with
the increase of seasonality that emphasized the cyclical
character of the siliciclastic-carbonate unit. An increase
of environmental instability led to small carbonate bodies
with a high percentage of siliciclastic components,
indicating only short-term and spatially restricted conditions
conducive to carbonate production.
2.	 Orbitally-forced cyclicity caused the alternating deposition
of carbonates and siliciclastics. The production
of carbonates was connected with dry periods that
caused a decrease of nutrient input from the continent,
oligotrophic conditions and the expansion of seagrass
meadows. The postulated seasonality probably led
to downwelling circulations with mixed warm oligotrophic
water masses in summer and a well-stratified
water column in winter. In the upper layer of the water
column, salinity fluctuations occurred. Siliciclastics
were deposited during a wet period with a concomitant
increase of nutrient input from the continent.
3.	 Carbonate bodies were deposited in various positions
within a paleodepth of the first tens of metres, whereby
accumulation of allochthonous carbonate material cannot
be excluded. They correlate with lowstand conditions.
Exceptionally, larger limestone bodies were
deposited under transgressive, highstand and lowstand
conditions. Differences in the individual carbonate
bodies reflect variations in the intensity of terrigenous
input, the dynamics of the environment, the depth
(above and below the fair-weather wave-base) and the
morphology of the sea-floor.
4.	 Siliciclastic intercalations in the limestone bodies
represent catastrophic rainfall events that transported
a higher amount of terrigenous material to the basin,
including phytodetritus, which represents a food
source for suspension-feeders. The input of freshwater
is responsible for a decrease in salinity.
Acknowledgments  This research was supported by grant nos.
PRVOUK P44 and GAČR 205/09/0103. We are grateful to Mrs. Sarka
Rousava for correction of the English language. The constructive
reviews of Natália Hudácˇková (Comenius University in Bratislava,
Slovakia) and of an anonymous reviewer substantially improved the
manuscript.
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GEOLOGICA CARPATHICA, JUNE 2011, 62, 3, 251—266 doi: 10.2478/v10096-011-0020-0
Miocene vegetation pattern and climate change in the
northwestern Central Paratethys domain
(Czech and Slovak Republic)
MARIANNA KOVÁČOVÁ1
, NELA DOLÁKOVÁ2
and MICHAL KOVÁČ1
1
Department of Geology and Paleontology, Faculty of Sciences, Comenius University, Mlynská dolina, 842 15 Bratislava,
Slovak Republic; kovacova@fns.uniba.sk; kovac@fns.uniba.sk
2
Institute of Geological Sciences, Masaryk University, Kotlářská 2, 611 37 Brno, Czech Republic; nela@sci.muni.cz
(Manuscript received October 13, 2010; accepted in revised form December 16, 2010)
Abstract: The case study area covers the slopes of the tectonically quiet European platform and foreland of the tectonically
active Carpathian mountain chain (Carpathian Foredeep and Vienna Basin). Therefore the research on pollen spectra
mirrors not only the evolution of landscape in two areas with different geodynamics, but also climatic changes in the
Central Paratethys domain during the studied time interval. According to the pollen data, the Early to Middle Miocene
vegetation reflects subtropical climate with very mild (negligible) cooling events during this period. This is indicated by
common occurrence of thermophilous taxa in the whole sedimentary record. The Middle Miocene landscape evolution,
conditioned by uplift of the Carpathian mountain chain and subsidence of adjacent lowlands, led to commencement of the
altitudinal zonation. The terrestrial and aquatic ecosystems confirm a subtropical climate (Miocene Climatic Optimum,
Mi3 event) with some possible long term changes in humidity. The Late Miocene paleogeographical changes, but also
general climatic oscillations in the northwestern Central Paratethys realm, resulted in decrease of the number of thermophilous
taxa during this time (change in latitudinal position of the vegetation cover). Variously high mountain relief of
the uplifted mountain chains (altitudinal zonality) created ideal conditions for mixed mesophytic forests (to open woodland
– open grassland type), still with presence of evergreen taxa. A subtropical climate with gradual transition to
warm temperate climatic conditions is supposed on the basis of the reconstructed vegetation cover.
Key words: Miocene, Paratethys, Carpathian Foredeep, Vienna Basin, paleoclimate, palynology.
Introduction
The Miocene vegetation pattern and climatic changes were
studied by means of palynology in the northwestern part of the
Central Paratethys domain (Fig. 1). To determine changes in
vegetation pattern (altitudinal zonation) and influence of the
global climatic changes (latitudinal zonation) two areas with
different geodynamics and therefore also with different landscape
evolutions have been choosen. To the West there was
the tectonically quiet Variscan Bohemian Massif, and to the
East the neo-Alpine tectonically active uplifting Western Carpathian
mountain chain. The samples were taken from marine,
brackish to freshwater sediments of the Carpathian Foredeep
(Czech Republic) and Vienna Basin (Czech and Slovak Republic)
in the time interval: Early to Late Miocene—Eggenburgian
to Pannonian (Burdigalian to Tortonian, sensu
Harzhauser & Piller 2007). The analysed sediments were well
biostratigraphically dated (Hladilová 1988; Nehyba et al.
1997; Doláková et al. 1999; Rögl et al. 2003; Hudáčková et al.
2003; Kováč et al. 2004, 2006, 2007, 2008).
Paleogeography
The study area – the contact zone between the North European
Platform and Western Carpathian orogen had a very
complicated Neogene geodynamic history (e.g. Royden
1985, 1988; Ratschbacher et al. 1991a,b; Kováč & Hók
1993; Lankreijer et al. 1995; Meulenkamp et al. 1996; Nehyba
et al. 1997; Kováč et al. 1997, 1998a,b, 2001, 2003; Kováč
2000; Kvaček et al. 2006; Harzhauser & Piller 2007). All
processes such as subduction, collision, back arc basin development
and its tectonic inversion are recorded in great paleogeographical
changes (Tomek & Hall 1993; Konečný et
al. 2002). These changes strongly influenced not only the relation
between areas flooded by the sea and continental areas,
but also development of landscape and the evolution of
altitudinal zonation in this region (Fig. 2).
The Early Miocene geodynamic development of the Bohemian
Massif and Western Carpathians junction area was
strongly affected by subduction of the flysch troughs basement
below the East Alpine—Western Carpathian orogen and
development of the Outer Western Carpathian accretionary
wedge from Flysch Belt units (Kováč et al. 1997, 1998a;
Kováč 2000; Konečný et al. 2002; Kvaček et al. 2006). Territory
in-between the platform and the front of the Carpathian
orogen was covered by a wide arm of the Central
Paratethys Sea during this time. The Eggenburgian marine
transgression flooded the Carpathian Foredeep on the slopes
of the Bohemian Massif in the West, continuing eastwards to
the Pouzdřany and Ždánice residual flysch troughs. The eastern
margin of the Early Miocene sea arm was represented by
the western and northern parts of the present Vienna Basin
(Fig. 2A). This sea arm represented a system of particular
www.geologicacarpathica.sk
252 KOVÁČOVÁ, DOLÁKOVÁ and KOVÁČ
GEOLOGICA CARPATHICA, 2011,62, 3, 251—266
basins with highly complicated shoreline contours and a
variable mutual communication with the open sea. As well
as a shallow marine environment with common lagoons and
deltas at the sea margin (Nehyba et al. 1997; Kováč et al.
1998a,b, 2004), a bathyal to neritic deep water sedimentary
environment with proved upwelling in the axial part of the
basin was documented (residual flysch troughs), still during
the Ottnangian (Roetzel et al. 2006; Grünert et al. 2010). The
Early Miocene transgression, following sea connection with
the Mediterranean via the Alpine Foredeep, can be correlated
Fig. 1. Geographical position of studied floral sites.
MIOCENE VEGETATION AND CLIMATE CHANGE IN CENTRAL PARATETHYS (CZECH AND SLOVAK REPUBLIC) 253
GEOLOGICA CARPATHICA, 2011,62, 3, 251—266
Fig. 2. Paleogeography of the Central Paratethys (according to Kováč 2000).
with the first Burdigalian global sea-level rise (sensu Vail et
al. 1977; Haq et al. 1988; Haq 1991; Kováč 2000).
At the end of the Ottnangian, the transpressive tectonics
resulted in a partial uplift of the Alpine-Carpathian chain,
followed by closing of the sea connections with the Mediterranean
in front of the Alps. Short term isolation of the Western
Carpathian sedimentary basins took place. This process
was accompanied by closing of the residual flysch troughs
and folding and thrusting of their sedimentary fill towards
the platform. Gradual uplift of the Outer Western Carpathian
accretion wedge started (Kováč et al. 1998a). On the other
side, initial rifting of the Pannonian back arc basin began in
the Western Carpathians hinterland (Horváth et al. 1988).
The Karpatian transpressive tectonics led to extrusion of
the Western Carpathian orogene (ALCAPA Microplate)
from the Alpine domain (Ratschbacher et al. 1991a,b; Kováč
et al. 1998a). Oblique collision between the orogen and the
Bohemian Massif resulted in left lateral displacement along
this zone followed by opening of the Vienna Basin “pull
apart” depocentres (Royden 1985; Tomek & Hall 1993;
Lankreijer et al. 1995; Fodor 1995; Kováč et al. 2004). Initial
rifting in the Pannonian back arc basin opened new marine
connections in this time. The sea transgression from the
Mediterranean advanced via the Trans-Tethydian Trench
Corridor (Rögl 1998; Pavelič 2001). The broad Karpatian
sea flooding extended beside the back arc basin also into the
Vienna Basin and the Western Carpathian Foredeep domain
(Fig. 2B). The continental areas were represented beside the
European platform by uplifted parts of internal zones of the
Eastern Alps and Western Carpathians; the accretion wedge
of the Outer Carpathian Flysch Belt showed a minimal uplift
in its western part only.
The Middle Miocene geodynamic development influenced
factors such as the end of subduction in front of the Western
Carpathian orogen, its soft docking on the slopes of the
European platform (Konečný et al. 2002) and back arc synrift
subsidence in the Pannonian Basin domain (Horváth et
al. 1988). Voluminous acid and calc-alkaline volcanism is
254 KOVÁČOVÁ, DOLÁKOVÁ and KOVÁČ
GEOLOGICA CARPATHICA, 2011,62, 3, 251—266
observed during this time. The Central Paratethys Sea can be
characterized in this time as an epicontinental sea with a
number of archipelagoes and a lot of separate basins. The sea
flooding extended in front of the orogen (foredeep basin), as
in the Pannonian back arc basin area. In the study area
(southern part of the Carpathian Foredeep and Vienna Basin),
the late Early Badenian transgression was completely
controlled by tectonics (Lankreijer et al. 1995; Kováč et al.
2001, 2004) and associated with basin subsidence and continuous
mountain chain uplift. During the Middle Badenian a
part of the Western Carpathian basins were isolated with the
resulting salinity crisis (Kováč et al. 1998b); in the southern
part of the Carpathian Foredeep the sedimentation ended.
The Vienna Basin still subsided and was connected towards
the Pannonian domain by straits in the uplifting Malé Karpaty
and Leitha Mountains. The changes in relief led to development
of a drainage system, the paleo-Danube river
delta entered the Vienna Basin and voluminous deltaic sequences
started to be deposited (Kováč et al. 2004; Lambert
et al. 2008). The Late Badenian flooding covered the whole
northen part of the Pannonian back arc basin system
(Fig. 2C). In the Vienna Basin the transgression was accelerated
by basin subsidence and sea-level rise (Kováč et al. 2001,
2006). The basins were filled by clastic material transported
by rivers, entering the basin from elevated mountain ranges in
the basin surroundings. In front of the orogene, the Carpathian
Foredeep started to disintegrate and depocentres moved from
West towards East (Meulenkamp et al. 1996). During the Sarmatian,
closing of the Central Paratethys Sea connections with
the Mediterranean Sea led to isolation and salinity decrease in
all the Western Carpathian basins. The sea started to be shallower,
but its extent did not change significantly.
The Late Miocene geodynamic development represents the
final stage of the Pannonian back arc basin evolution, with related
thermal subsidence (Horváth 1988; Kováč et al. 1993;
Lankreijer et al. 1995; Konečný et al. 2002). The Vienna Basin
represented a partly isolated bay at its northwestern boundary
during this time (Kováč et al. 1998a; Kvaček et al. 2006).
Uplift of the Western Carpathian mountain chain was accompanied
by the next development of the river net, resembling
the Pliocene paleogeography (Fig. 2D). The Central Paratethys
brackish sedimentary environment – represented by
Lake Pannon in the study area (Magyar et al. 1999) gradually
changed into a freshwater lake environment, particularly in
the back arc basin domain. During the Pannonian, the Vienna
Basin was filled by a huge amount of deltaic sediments
(Kováč et al. 1998, 2004, 2006; Harzhauser et al. 2004). The
shallow-water fluvial to lacustrine environment changed to
swamps and alluvial plains with ephemeral lakes representing
the greater part of its territory until the end of the Late Miocene.
The Pannonian and Pontian mountain ranges gained features
similar to their present form.
Material and methods
In this study, 44 outcrops and boreholes (Carpathian Foredeep
– Eggenburgian—Early Badenian (23 localities), Czech
andSlovakparts of Vienna Basin– Karpatian—Pannonian
(21 localities)) were analysed. Due to the absence of index
fossils, the studied Lower Miocene sediments are undistinguishably
(Upper Eggenburgian—Ottnangian). The analysed
samples come from the localities in the southern part of the
Carpathian Foredeep – boreholes Šafov 12, Šafov 13, Čejkovice,
Únanov, Miroslav, Trboušany, Nosislav 3, Židenice.
Carpathian Foredeep marine and brackish sediments, Karpatian
in age, were evaluated from several stratotype localities
Slup, Hevlín, Dolní Dunajovice, Medlov, boreholes Nosislav
3, Ždánice 67, 68 and from the Vienna Basin boreholes
Zohor 1 and Gbely 139 were analysed (Doláková &
Slamková 2003). Palynological data, Early Badenian in age,
come from the Carpathian Foredeep marine sediments from
the localities Židlochovice, Lysice, boreholes Moravské
Knínice, Sivice, Chrlice, Opatovice and Otmarov. Pollen
data, Late Badenian in age, come from outcrop Devínska
Nová Ves and borehole Lozorno 1 in the Vienna Basin. Studied
sediments, Late Miocene in age, with well-determined
plant macrofossils (Knobloch 1968, 1985) come from the Poštorná,
Dubňany, Moravská Nová Ves outcrops; clay pit Gbely,
boreholes Suchohrad 32, Suchohrad 38, Jakubov 54 and six
shallow Pohansko boreholes near Břeclav city.
In the chemical treatment 20—30 g of dry sediment was
used. The samples were treated with cold HCl (35%) and HF
(70%), removing carbonates and silica. Separation of the palynomorphs
from the rest of the residue was carried out using
ZnCl2 (density=2 g/cm3
). Sieving was done using 10 µm nylon
sieve. The palynological residue, mixed with glycerine,
was prepared on slides. A transmitted light microscope with
250, 400, 630, 1000 (oil immersion) magnifications and SEM
microscope was used for pollen counting and identification.
Original micrographs are housed at the Institute of Geological
Sciences MU in Brno. The pollen diagrams have been created
using POLPAL 4 software (Walanus & Nalepka 1999).
To have a better idea about the vegetation composition the
differentiated vegetation groups (zonal, azonal, extrazonal)
were used sensu Kovar Eder et al. (2008a,b) and Kvaček et
al. (2006). A semiquantitave evaluation of climate evolution
has been done based on the proportion of paleotropical (thermophilous)
and arctotertiary elements (sensu Mai 1981,
1991) in terms of mesophytic plants. We devided the floristic
elements of zonal vegetation into two groups.
In the thermophilous-mesophytic group we included Engelhardia,
Sapotaceae, Palmae, evergreen Fagaceae (including
morphospecies Quercoidites microhenrici and Quercoidites
henrici), Trigonobalanopsis, Symplocos, Cornaceaepollis
satzveyensis, Tricolpopollenites liblarensis, Araliaceae, Rutaceae.
Mostly broad-leaved deciduous elements of warm-temperate
mixed mesophytic forests such as Quercus, Celtis,
Carya, Tilia, Zelkova, Ostrya, Carpinus, Betula, Juglans are
included into the group of arctotertiary—mesophytic elements.
Extrazonal mountain vegetation is represented by: Cedrus,
Tsuga, Picea, Cathaya. Azonal vegetation is influenced by
edaphic factors and in our study it is represented by riparian
forests with Alnus, Salix, Ulmus, swamps with Taxodiaceae,
Myricaceae, Nyssaceae and aquatic plant communities.
All the studied material is housed at the Institute of Geological
Sciences MU in Brno and the Faculty of Sciences of
Comenius University in Bratislava.
MIOCENE VEGETATION AND CLIMATE CHANGE IN CENTRAL PARATETHYS (CZECH AND SLOVAK REPUBLIC) 255
GEOLOGICA CARPATHICA, 2011,62, 3, 251—266
Early Miocene
Results and discussion During the Karpatian the thermophilous elements like Rutaceae,
Symplocos and Platanus occurred less regularly. Temperate
taxa are represented generally in low frequencies, but
the amount and diversity of the temperate mesophytic eleEggenburgian—Ottnangian—Karpatian
(Late Aquitanian—
Late Burdigalian)
During the Early Miocene thermophilous taxa Engelhardia,
Platycarya, Sapotaceae, Palmae and ferns Lygodium, Pteridaceae,
?Davalliaceae, Schizaeaceae—Cyatheaceae were frequent.
Evergreen Fagaceae were represented by the
Trigonobalanopsis type, morphotaxa Tricolporopollenites
microhenrici and Tricolpopollenites liblarensis, Tricolporopollenites
henrici. Also Symplocos, Reevesia, Parthenocissus,
Araliaceae, Rutaceae and morphotaxa Cornaceaepollis
satzveyensis, Tricolporopollenites pseudocingulum were common
in pollen spectra. Arctotertiary elements Carya, Juglans,
Quercus, Betula, Liquidambar were less frequent (Fig. 3).
The vegetation of the salt marshes and also insolated places
(Chenopodiaceae up to 37 %, Ilex, Tamarix, Ericaceae, Poaceae
less Ephedra, Asteraceae and Buxaceae) was typical. Due to
the salinity oscillations as well as occasional higher evaporation
sensu Hladilová (1988), the coasts of individual sea gulfs
and lagoons could be repeatedly salinized and overgrown by
the halophilous flora (Doláková et al. 1999). Pollen grains of
the formal genus Monocirculipollis assigned to the family
Caryophyllaceae (sensu Doláková 2004), were typical for this
time span being absent in younger ones (Fig. 6). Salt marsh
vegetation was sometimes replaced by swamp plants.
Taxodiaceae, Myricaceae, Cyrillaceae, Gleicheniaceae,
Decodon, Lygodium, Selaginella. Even the aquatic flora appeared
– Sparganium, Potamogeton, Onagraceae, Nelumbo,
Cyperaceae (Figs. 3, 5, 7). The genus Platanus was the common
member of the pollen spectra since this time span. A permanently
low amount of intrazonal elements (Taxodiaceae,
Myricaceae and ferns) without strong oscilation occurred in the
palynospectra. Regularly higher ratios of Fagaceae, Carya and
also heliophilous taxa were observed. Pinaceae (up to 40 %)
and extrazonal vegetation, including abundant Cathaya and less
frequent Cedrus, Picea, Abies, occurred frequently (Fig. 3). A
high proportion of Ulmaceae, Myrica, Alnus was observed in
sediments of the Eggenburgian—Ottnangian. Pollen spectra contain
a larger amount of spores of thermophilous ferns as Lygodium
(up to 5 %), Pteridaceae, Gleicheniaceae together with
Selaginella and bryophyte Riccia (Fig. 3). These findings are in
a good conformity with macrofloristic results of Knobloch
(1982) who described a unique oryctocoenosis from the rhyolite
tuffites at Znojmo and Přímětice. He considered them as the
shrubby – arboreal heliophilous vegetation with mostly evergreen
fine dentate or spiny leave (sclerophyllous) similar to
Mediterranean “macchias”. Swamp vegetation with Glyptostrobus,
Myrica, aquatic flora with Salvinia, Potamogeton, Nymphaea
and coastal reed with Typha, Decodon, Sparganium were
identified based on macrofloristic remains too (Knobloch
1982). The accumulations of Limnocarpus fruits growing in the
brackish water were described by Knobloch (1984). The pollen
often found in clumps (Myricaceae, Chenopodiaceae, Caryophyllaceae,
Oleaceae, Onagraceae, Platanus) support the low
water dynamics and a short transport (Figs. 5, 6, 7).
ments slightly increased (Fig. 4). Mountain vegetation with
Tsuga and Abies was common (Fig. 4). Subtropical humid climate
was also supported by the macrofloristic remains described
from the stratotype localities Slup and Dolní
Dunajovice. Leaves of the family Lauraceae predominated
with small proportion of deciduous trees in this association
(Knobloch 1967, 1982; Kvaček 2003). The azonal vegetation
is represented by swamp and riparian forests dominated by
Glyptostrobus and Myrica.
Marshy-palm forest with Calamus, Poaceae, Lygodium,
Sparganium, Potamogeton and riparian forest with Alnus,
Ulmus, Myricaceae, Lythraceae or Selaginella are frequent
(Figs. 4, 8). The associations with Taxodiaceae, Craigia and
Pteridaceae (up to 10 %) and Polypodiaceae document a well
developed swamp environment (Figs. 4, 8). During the
present time the genus Craigia occurs in broad-leaved evergreen
and deciduous mixed forests and seasonally wet forests.
However, according to Kvaček et al. (2002), ecological
tolerances of its fossil representatives may have been greater
during the Tertiary. This tree surely tolerated swampy conditions
and entered even coal-forming forests in wetland habitats
namely swamp forests dominated by the Taxodiaceae
and many other swampy and riparian woody plants as well
aquatic herbs. Konzalová (1976) described a very similar horizon
with Intratriporopollenites insculptus Mai (Craigia)
from the coal seam formation and Cypris claystones of the
North Bohemian basins.
During this time interval the plant assemblages with higher
portion of arctotertiary elements were described by several
authors from the Silesian part of the Carpathian Foredeep in
the Polish Lowland (Oszast & Stuchlik 1977; Stuchlik 1980;
Sadowska 1989; Ważyńska et al. 1998). The decrease of
thermophilous elements during the Ottnangian—Early Karpatian
has been recognized and defined as the microfloristic
Zone MF-3 by Planderová (1990) and Planderová et al.
(1993a,b). Such an event has never been found in the Carpathian
Foredeep. This fact is probably related to different
paleogeography. The most similar pollenspectra of Karpatian
age was published by Nagy (1999) from the Mecsek Mts.
Environmental interpretation of data from the Carpathian
Foredeep is similar to the conditions in the Korneuburg Basin
(Hofmann et al. 2002), except for absence of Avicenia and
the lower portion of the Palmae in the studied area. Early
Miocene was the warmest period of the Miocene in the Pannonian
Basin and the sporomorphs indicate a warm subtropical
climate (Nagy 2005).
Middle Miocene
Badenian—Sarmatian (Langhian—Serravallian)
Swamp elements (Taxodiaceae) have more regular occurrence
without oscillations in comparison with the Karpatian.
Olea type pollen was less frequent in comparison with Lower
Miocene pollen spectra. In the Badenian pollen spectra the
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GEOLOGICA CARPATHICA, 2011, 62,3, 251-266
258 KOVÁČOVÁ, DOLÁKOVÁ and KOVÁČ
GEOLOGICA CARPATHICA, 2011,62, 3, 251—266
Fig.5.EarlyMiocenesporomorphsfromthestudiedarea.1–Chenopodiaceaeaccumulation,2–Asteraceae—Asteroideae–TricolporopollenitesgrandisNagy,3–Asteraceae—Cichorioideae–CichoreaciditesgracilisNagy,4–Oleaceae–(Oleoidearumpollenitessp.),5–
Oleaceae
accumulation, 6 – Ephedra sp. + Chenopodiaceae, 7 – Ericaceae (Ericipites callidus (Potonié) Krutzsch), 8 – Caryophyllaceae (Monocirculipollis
sp. Krutzsch), 9 – Caryophyllaceae accumulation, 10 – Mastixia – (Tricolporopollenites satzvayensis Pflug), 11 – Myrica accumulation,
12 – Taxodiaceae – ?Glyptostrobus sp., 13 – Potamogeton – (Potamogetonicidites paluster (Mamten) Mohr), 14 – Nelumbo sp.
(Nelumbopolleniteseuropaeus(Tarasewich)Skawińska),15–Cyperaceae–(CyperaceaepollispiriformisThile-Pfeifer).
MIOCENE VEGETATION AND CLIMATE CHANGE IN CENTRAL PARATETHYS (CZECH AND SLOVAK REPUBLIC) 259
GEOLOGICA CARPATHICA, 2011,62, 3, 251—266
Fig. 6. Eggenburgian palynomorphs. 1a,b, 3, 4a,b – Tamarix sp.; 1, 3, 4 – LM 1000ξ; 1a – SEM 5500ξ; 1b – SEM 12,000ξ; 4a –
SEM 5500ξ; 4b – SEM 12,000ξ. 2 – Tamarix gallica – recent – LM 1000ξ; 2a – SEM 6000ξ. 5a,b, 6 – Salix sp.; 5 – LM 1000ξ;
5a – SEM 5000ξ; 5b – SEM 12,000ξ. 6 – SEM 6000ξ. 7a,b, 8, 9 – Rutaceae; 7, 8, 9 – LM 1000ξ; 7a – SEM 5000ξ; 7b –
SEM 12,000ξ. 10, 11,
12 – Platanussp.;10a,b, 11, 12 – LM 1000ξ; 10a – SEM 4000ξ; 10b – SEM 10,000ξ.
260 KOVÁČOVÁ, DOLÁKOVÁ and KOVÁČ
GEOLOGICA CARPATHICA, 2011,62, 3, 251—266
Fig.7.Karpatiansporomorphsfromthestudiedarea.1–Engelhardiasp.withcavitiesafterpyritecrystals;2–Palmae–Monocolpopolle-
nitestranquillus(Potonié)Thomson&Pflug;3–Palmae–Arecipitessp.;4–Poaceae–Graminiditessp.;5–Sparganiumsp.–(Sparganiaceaepollenites
neogenicus Krutzsch); 6 – Alnus sp. – (Alnipollenites verus (Potonié) Potonié); 7 – Myrica sp. (Myricipites coryphaeus
(Potonié)Potonié);8–Myricasp.(Myricipitesperegriniformis(Gladkova)Grabowska&Wazynska);9,10–Lythraceae;11,12–Craigia
sp. (Intratriporopollenites insculptus Mai); 13 – Taxodiaceae – ?Glyptostrobus sp.; 14 – Selaginella sp. (Echinatisporis miocenicus
Krutzsch & Sontag in Krutzsch); 15 – Pteris sp. – (Polypodiaceoisporites muricinguliformis Nagy); 16 – Ilex sp. – (Ilexpollenites
margaritatus(Potonié) Raatz);17–Tsugasp. –(Tsugaepollenitesmaximus (Raatz) Nagy; 18–Lygodium sp. (Leiotriletesmaxoides
maxoides W.Kr.).
MIOCENE VEGETATION AND CLIMATE CHANGE IN CENTRAL PARATETHYS (CZECH AND SLOVAK REPUBLIC) 261
GEOLOGICA CARPATHICA, 2011,62, 3, 251—266
Fig.8.Badeniansporomorphsfromthestudiedarea.1–Mastixiasp.–(TricolporopollenitessatzvayensisPflug);2–Sapotaceae–(Sapotaceoipollenites
sapotoides (Pflug & Thomson) Potonié); 3 – Quercoidites henrici (Potonié) Potonié, Thomson & Thiergart; 4, 5 – Quercoidites
microhenrici (Potonié) Potonié, Thomson & Thiergart; 6, 7 – Q. robur (Quercoidites granulatus (Nagy) Slodkowska); 8 – Quercoidites sp.;
9–Zelkovasp.–(ZelkovaepollenitespotonieiNagy),10–Tiliasp.—(Intratriporopollenitesinstructus(Potonié)Thomson),11–Loran-
thaceae–GothanipollenitesgothaniKrutzsch;12–Symplocossp.–Symplocoipollenitesvestibulum(Potonié)Potonié,13–Cercidiphyllum
sp.–(Cercidiphyllitesminimireticulatus(Trevisan)Ziembińska-Tworzydło);14,15–Distylium–Parrotiatype–(Tricolporopollenitesindeterminatus(Romanovicz)
Ziembińska-Tworzydło);16 – Platanussp.;17 –Pteridaceae –(Segmentizonosporitespaucirugosus(Nagy)
Stuchlik); 18 – Caryophyllaceae (Caryophyllidites microreticulatus Nagy); 19 – Marine dinoflagellates; 20 – foraminiferal lining.
262 KOVÁČOVÁ, DOLÁKOVÁ and KOVÁČ
GEOLOGICA CARPATHICA, 2011,62, 3, 251—266
higher differentiation of the Fagaceae in thermophilous evergreen
(morphotypes Tricolporopollenites henrici and Tricolporopollenites
microhenrici) and deciduous oaks, some
thermophilous taxa Gothanipollenites gothani (Loranthaceae)
or Tricolporopollenites indeterminatus (Hammamelidaceae)
occurred (Fig. 9). Herbs such as Caryophyllaceae (Minutipollis
granulatus Krutzsch) were common. Early Badenian Lauraceae
and Betulaceae leaves from the Carpathian Foredeep
have been found at the Smolín locality (Sitár et al. 1978). During
the Late Badenian the following were frequently present:
Pinaceae (Pinus, Picea, Abies, Tsuga) and deciduous elements
with Quercus, Alnus, Ulmus, Carya. Subtropical taxa are commonly
represented by Magnolia, Platycarya, Engelhardia,
Myrica, Trigonobalanopsis and Distylium. Paleoecological
conditions favoured development of swamp forests with Taxodiaceae,
Nyssa and Myrica and riparian forest elements with
Alnus, Ulmus and Pterocarya. Drier areas were overgrown
with mixed mesophytic forest represented by Pinus, Juglans,
Carya, Sciadopitys and the extrazonal vegetation type is documented
by the presence of Picea, Tsuga and deciduous oaks.
Herbs are represented mostly by Poaceae. During the Early
Sarmatian time interval the azonal vegetation was well developed
in swamps with Taxodiaceae, Myrica, Nyssa and salty
marshes with Poaceae and halophytes (Chenopodiaceae).
Riparian forests with Alnus, Salix and Ulmus were also common.
The extrazonal vegetation portion increased mainly in
the mountain vegetation elements. The decrease to disappearance
of the Taxodiaceae, Nyssaceae, Myricaceae and halophytes
suggests a large reduction of the swamp biotops.
Riparian forests with Ulmus, Salix, Alnus and Poaceae were
still present. Gradual decrease of thermophilous taxa indicates
moderate cooling. Syabryaj & Vodoryan (1975) described similar,
well diversified pollen spectra from the NE Carpathian
territory in Čop—Munkacevo. Ivanov (1995) and Ivanov et al.
(2002) described presence of Symplocos in Badenian pollen
spectra from NW Bulgaria. In our studied material we noticed
Symplocos presence only from Early Miocene localities, probably
due to temperature gradient.
There was a warm subtropical climate. Early Badenian
transgression and uplift of the Alpine and Carpathian Mountains
produced favourable local climate for vegetation
change (Nagy 2005).
Late Miocene
Pannonian—Pontian (Tortonian—Messinian)
In the pollen spectra from the Early Pannonian sensu
Harzhauser et al. (2004) or Early Tortonian sensu Harzhauser &
Piller (2007), mostly broad-leaved deciduous elements dominate,
with some thermophilous elements admixture of Engelhardia,
Ilex, Castanopsis and Castanea, suggesting a warm
temperate mixed mesophytic forest with low representation of
evergreen elements. The proportion of NAP – non arboreal
pollen – Ericaceae and Chenopodiaceae is higher (10 and
14 % respectively) suggesting local marshes and open herbaceous
plant communities within the forests. Mountain conifers,
such as Picea, Tsuga, Abies, Cedrus are common accessories.
Lowland vegetation was comprised of the azonal Alnus, Pinus,
Ulmus mixed and broad-leaved riparian forest with common
deciduous oaks, and swamp taxa Taxodiaceae, Nyssa, Myrica.
Sporadic occurrences of dinoflagellates and green algae Tasmanaceae
indicate a slightly higher salinity, Botryococcus can
thrive in both brackish or freshwater environments, whereas
green algae Pediastrum, Mougeotia, aquatic ferns Azolla, and
aquatic and coastal plants (Nelumbo, Nymphaea, Myriophyllum,
Sparganium, Potamogeton etc.) represent a freshwater environment
(Doláková & Kováčová 2008). In the pollen spectra,
from the middle Pannonian (sensu Harzhauser et al. 2004), coniferous
woody plants of mountain vegetation (Picea, Abies,
Tsuga, Cedrus, Pinus) and deciduous oaks were abundant. Angiosperm
trees and shrubs with Alnus, Betula, Liquidambar,
Myrica, Nyssa and Salix indicate a well developed riparian forest.
The subdominance of herb species is good evidence of the
local open woodland environment.
The facies mutually changing in time and space in individual
pollen spectra are created by azonal types of vegetation
(marshes, riparian, coastal and aquatic) or by high amounts
of herbaceous plants Artemisia, Plantago, Polygonum,
Asteraceae, Lamiaceae, Daucaceae, Caryophyllaceae, which
indicate existence of local open areas.
In the Slovak part of the Danube Basin Planderová (1972,
1990) described reduced marshes, isolated lakes surrounded
by steppe meadows (dominance of Artemisia) with rare
woody plants. In comparison with Hungary she considered
the climate cooler and drier (Nagy 1985; Nagy & Planderová
1985; Planderová 1990). Hoffmann & Zetter (2005) determined
a pollen assemblage rich in herbs from the Late Pannonian
in the Styrian Basin.
The extensive Pannonian and Pontian sea and the protective
mountain range provided a very equable, warm temperate
climate, where even the summer season was not too dry
(Nagy 2005).
Conclusions
Development of Miocene vegetation patterns in the area of
the northwestern Central Paratethys was derived (above all)
from palynological analysis. The case study area covers the
slopes of the tectonically quiet European platform and the
foreland of the tectonically active Carpathian mountain chain
(Carpathian Foredeep and Vienna Basin). Interpretation of
pollen spectra reflects both, the landscape evolution in two
areas with different geodynamics, and the climatic changes in
the Central Paratethys domain during the studied time intervals.
Based on pollen data, the Early to Middle Miocene vegetation
document a subtropical climate with very mild
(negligible) cooling during this period. This is indicated by
common occurrence of thermophilous taxa: Sapotaceae,
Palmae, Engelhardia, Platycarya and Tricolporopollenites
henrici, Lygodium and Pteridaceae. Reevesia, Cornus-Mastixia,
Symplocos, Parthenocissus, Tricolporopollenites pseudocingulum,
Rutaceae and Araliaceae. The proportion of the temperate
elements such us Carya, Pterocarya, Juglans, Celtis,
Fagus is noticeably lower. The lower portion of extrazonal
(mountain) vegetation and well developed riparian forests
with Alnus, Craigia, Pteridaceae, Polypodiaceae, Lythraceae,
MIOCENE VEGETATION AND CLIMATE CHANGE IN CENTRAL PARATETHYS (CZECH AND SLOVAK REPUBLIC) 263
GEOLOGICA CARPATHICA, 2011,62, 3, 251—266
Fig.9. Pannonian palynomorphs from the studied area. 1 – Picea sp. – (Piceapollis sp.); 2 – Nyssa sp. – (Nyssapollenites rodderensis
(Thiergart) Kedves);3 –Quercusroburtype;4 –Salixipollenites sp.+Alnipollenitessp.;5 –Rosaceaegen.indet.;6,7 –Artemisia div.
sp.;8–Cichorioideae–(CichoreaciditesgracilisNagy);9–Asteroideae—(Tubulifloriditesmacroechinatus(Trevisan)Nagy);10–Centaureajaceatype;11–Daucaceaegen.indet.;12–Caryophyllaceaegen.indet.;13–Polygonumpersicaria–(Persicarioipollis
pliocenicus
Krutzsch); 14 – Pediastrum simplex Meyen; 15 – Microsporangium with glochidium of Azolla bohemica Pacltová.
264 KOVÁČOVÁ, DOLÁKOVÁ and KOVÁČ
GEOLOGICA CARPATHICA, 2011,62, 3, 251—266
Cyperaceae, Sparganium, Potamogeton, Nelumbo and swamps
with Taxodiaceae, Myricaceae alternated with salt marshes
represented by Chenopodiaceae up to 37 %, Poaceae, Caryophyllaceae,
Asteraceae, Ericaeae, Lythraceae, which document
a moderate relief of landscape during the whole Early
Miocene. The frequent pollen clumps support a theory of the
low water dynamics and a short transport distance. The alteration
in palynomorphs caused by the crystallization of pyrite
in anoxic conditions was observed.
The Middle Miocene landscape evolution, conditioned by
the uplift of the Carpathian mountain chain and subsidence of
adjacent lowlands, led to commencement of the altitudinal zonation.
This process is documented by changes in paleovegetation
cover. In spite of this presence of zonal vegetation with
evergreen broadleaved forests supplemented by azonal vegetation
(riparian forests, swamps) is typical of the Early Badenian.
Only several thermophilous plants, such as Engelhardia
and Platycarya, which were frequent in all of the Early Miocene
associations, decreased in the Early Badenian pollenspectra.
From the Late Badenian a higher proportion of
extrazonal (mountain) vegetation were present in pollen spectra
(Picea, Abies, Tsuga, Cedrus). The terrestrial and aquatic
ecosystems confirm a subtropical climate with visible changes
at the boundary between the Early and Late Badenian. An increased
proportion of the arctotertiary taxa during the Late
Badenian is documented in pollenspectra by Quercus, Ulmus
and Carya, whereas Platycarya, Engelhardia, Myrica, Distylium
and thermophilous Fagaceae are less frequent. Herbs are represented
mainly by the halophytes (Chenopodiaceae).
Vegetation during the Early Sarmatian time interval was
formed by swamp elements with Taxodiaceae, Myricaceae,
Nyssaceae. High elevation species of woody plants Tsuga,
Picea, Cedrus, Abies are indicative of mountainous relief resulting
from volcanic activity. During the Late Sarmatian the
proportion of swamp elements decreased and was replaced
mostly by riparian forests.
The Late Miocene paleogeographical changes and general
climatic oscillations in the northwestern Central Paratethys
realm reflected the decrease especially of the thermophilous
taxa Engelhardia, Castanea, evergreen Fagaceae Quercoidites
microhenrici, and to a lesser extent of Quercoidites henrici,
Trigonobalanopsis, Symplocos, Cornaceaepollis satzveyensis,
Tricolpopollenites liblarensis. An apart from the mountain
vegetation the amount of herbaceous plants in the pollen spectra
increased during this time span. The varying height’s of the
moutain relief of the uplifted mountain chains (altitudinal
zonality) created ideal conditions for extrazonal vegetation
(Cedrus, Tsuga, Picea) and dominance of mixed mesophytic
forests with Quercus, Celtis, Carya, Tilia, Zelkova, Ostrya,
Liquidambar, Carpinus, Betula, Juglans and with regular
presence of evergreen taxa.
The swamp, riparian, often hydrophilous (Azolla, Nymphaea,
Potamogeton) and halophytic (Chenopodiaceae)
plants represent coastal swamps, local lagoons, and marshlands.
The higher percentage of the herbs (Artemisia, Asteraceae,
Lamiaceae, Polygonum, Daucaceae, Caryophyllaceae,
Plantago) and shrubs in the comparison with older time intervals,
shows that local open woodland – open grassland
started to develop during the Pannonian.
The gradual retreat of areas flooded by the sea, as well as
following retreat of the lake and swamp environment was
confirmed by the decrease of azonal vegetation towards the
end of this period. The reconstructed vegetation cover suggest
a subtropical climate with gradual transition to warm
temperate climatic conditions.
Acknowledgments: The authors wish to express their gratitude
to reviewers L. Hably, Z. Kvaček and D. Ivanov for
useful and constructive comments. This work was supported
by the Slovak Research and Development Agency under the
contracts APVV-LPP 0120-06, APVV-0280-07, ESF-EC-
0006-07, ESF-EC-0009-07 and the Slovak Grant Agency
VEGA (Projects No. 2/0060/09, 1/0483/10) and Czech Grant
Agency under the contract No. 205/09/0103.
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GEOLOGICA CARPATHICA, AUGUST 2006, 57, 4, 295—310
www.geologicacarpathica.sk
Miocene evolution of landscape and vegetation in the Central
Paratethys
ZLATKO KVAČEK1
, MICHAL KOVÁČ2
, JOHANNA KOVAR-EDER3
, NELA DOLÁKOVÁ4
, HENRIETTE
JECHOREK5
, VALENTIN PARASHIV6
, MARIANNA KOVÁČOVÁ2
and UBOMÍR SLIVA2
1
Charles University, Faculty of Science, Albertov 6, CZ-128 43 Praha 2, Czech Republic; kvacek@natur.cuni.cz
2
Department of Geology and Paleontology, Faculty of Sciences, Comenius University, Mlynská dolina, SK-842 15 Bratislava,
Slovak Republic; kovacm@fns.uniba.sk; kovacova@ fns.uniba.sk
3
Staatliches Museum für Naturkunde Stuttgart, Rosenstein 1, D-07191 Stuttgart, Germany; eder.smns@naturkundemuseum-bw.de
4
Department of Geology and Paleontology, Faculty of Sciences, Masaryk University, Kotlářská 2, CZ-611 37 Brno, Czech Republic;
nela@sci.muni.cz
5
Carl-von-Ossietzky-Straße 5, D-02826 Görlitz, Germany; hjechorekgr@surfeu.de
6
Faculté de Géologie, Université de Bucharest, Str. N. Balcescu, Bucharest, Romania; paleovaly@yahoo.com
(Manuscript received June 27, 2006; accepted in revised form December 8, 2005)
Abstract: The digital elevation model (DEM) helps to express Neogene landscapes and vegetation on palinspastic maps
with reconstructed orography. To reconstruct ancient vegetation cover, basic zonal vegetation formations and their characteristics
have been defined based on diversity and proportions of zonal woody evergreen, deciduous, sclerophyllous
and legume-type elements, besides intrazonal (azonal, e.g. coal-forming, aquatic and riparian) and extrazonal (montane
conifer-rich) vegetation. Three time intervals have been analysed – Karpatian to Early Badenian, Late Badenian to earliest
Sarmatian and Early to Middle Pannonian. After evaluating respective local sites of leaf, fruit/seed and spore/pollen
assemblages, paleogeobotanical maps have been constructed for the area of the Central Paratethys and its periphery.
Key words: Miocene, Central Paratethys, paleogeography, palinspastic maps, digital elevation models, vegetation mapping.
Introduction
It is a common task of geobotany today to express interpreted
vegetation over larger areas on maps, because the extent
of various types of plant communities is an important factor,
for example, in tracing human influence or migrations
of terrestrial animals. However, several aspects of such studies
are different considering Neogene vegetation (KovarEder
et al. submitted). The paleogeographic configuration
of land and sea was different from the present situation. The
regional relief changed in connection with orogeny processes.
The floristic spectra included elements mostly extinct
or no longer living in Europe. The time slices for the
respective maps are many million years distant from the Recent.
Global climate, atmospheric circulations and the
world ocean varied depending on the time interval studied.
To overcome these problems a team of specialists is needed.
The paleogeographic background with an approximate demarcation
of sea, basins and approximate relief is the first
premise to attempt such a paleogeobotanical mapping.
Complex and well determined spectra of plant elements
from the reference sites with both megafossil and spore/pollen
records are most relevant. The megafossil record usually
reflects the situation near the site and is differentiated according
to the lithofacies. It also reflects the presence of
plants producing poorly preservable pollen (e.g. Lauraceae)
and indicates more strongly floristic changes than the
spore/pollen spectra. Leaves may convey information on
the vegetation physiognomy. Fruits and seeds are better indicators
of systematic affinities. Pollen and spores may undergo
long distance transport by wind and their spectra thus
include information on the composition and changes of upland
vegetation, which usually consists of mountain conifer-rich
forests. It is problematic to transfer frequencies of
any kind of plant organs in the fossil spectra into true abundances
of plants in a community or landscape, because the
fossil record is biased either by overproduction of fossil organs
(e.g. pollen, diaspores) and taphonomic processes (e.g.
deciduous vs. evergreen foliage, more rapid decay of delicate
leaves). In our analyses, we relied mostly on qualitative
proportions of elements, that is the floral diversity, within
the given assemblage.
A synthetic view over a large area that includes several
countries should be obtained based both on own experiences
including authentic knowledge of plant fossil sites
and the review of the published data. In the latter cases, it
is often a difficult task to critically re-evaluate older taxonomical
interpretations, both in the megafossil and pollen
spectra. Particularly the interpretations of various pollen
types within the natural system look very different today
from the traditional morphological or semi-natural systems
used previously (or even at present). Particularly by
efforts of large-scale comparisons with living plants
(Stuchlik 1994) and electron scanning microscopy
(Walther & Zetter 1993; Ferguson et al. 1998; Zetter
1998; Liu et al. 2001), surprising solutions for several
sporomorphs have been suggested and important natural
affinities have been recognized (e.g. Mastixia, i.e. Cornaceaepollis
satzveynsis, Trigonobalanopsis, i.e. Castaneoipollenites
pusillus, etc.). Studies of pollen in situ and
296 KVAČEK et al.
co-occurrence with megafossils are also important. Thus it
has recently become obvious that the tilioid pollen does not
belong in most cases to Tilia, a deciduous zonal element, but
to Craigia & Dombeyopsis lobata or Banisteriaecarpum &
Byttneriophyllum tiliaefolium (alias “Alangium”) plants both
intrazonal, frequent constituents of the Glyptostrobus swamp
forests (e.g. Kvaček et al. 2002).
This paper introduces a new methodology of the Neogene
paleogeobotanical mapping presenting it on three
time slices of the Neogene in the Central Paratethys.
Methodology
Digital Elevation Models (DEMs) of the Central Paratethys
(Fig. 1) in individual time intervals of the Miocene,
which help us to express landscapes with expected orography,
were constructed on the basis of present knowledge
of geodynamic evolution of the Alpine-Carpathian-Pannonian
region (Csontos et al. 1992; Kováč et al. 1993,
1997, 2001, 2002; Meulenkamp et al. 1996; Baráth et al.
1997; Plašienka et al. 1997; Plašienka & Kováč 1999;
Bezák et al. 2002; Konečný et al. 2002; Soták & Kováč
2002; Bielik et al. 2004), existing palinspastic maps
(Kováč et al. 1989, 1998, 2003; Magyar et al. 1999;
Kováč 2000; Popov et al. 2004), as well as burial and uplift
history of sedimentary or other rock complexes (Horváth
et al. 1988; Dunkl 1992; Kováč et al. 1994; Hurai et
al. 1995; Dunkl & Demény 1997; Danišík et al. 2004,
etc.).
Models of the Central Paratethys vegetation (Figs. 2—4)
use a simplified system of vegetation units (formations),
which are usually sufficient to get tentative pictures of paleovegetation
in space. Our attention has been paid to distinguish
zonal, intrazonal (azonal) and extrazonal
formations on the basis of autecology and leaf physiognomy
of elements, whose grouping has been attempted in
this respect (Kovar-Eder & Kvaček 2003; Kovar-Eder et al.
submitted). The characteristics of the elements have been
mostly derived from autecologies of their nearest living
relatives or analogues. Reference fossil localities/plant assemblages
often included taxa of different vegetation formations,
of which zonal elements are relevant for the maps
of the reconstructed fossil vegetation. However, it is apparent
from the sedimentary settings that assemblages
dominated by intrazonal elements prevailed in the record
of megafossils, mainly from the basin deposits. Extrazonal
conifer-rich mountain vegetation was represented in pollen
spectra, exceptionally in megafossil records from the
intra-montane basins. Not only altitude, but also the direction
of exposure of mountain slopes and substrate may
have influenced the composition of the conifer stands.
Volcanic settings are the best environments to bring information
on zonal vegetation of mesic habitats. Percentages
of zonal herbs as well as Non Arboreal Pollen (NAP) as a
whole in pollen spectra may refer to close canopy forests
versus open woodland to steppe vegetation. Even intrazonal
elements can bring information on the character of
climate (e.g. the presence of palms), although the intrazonal
assemblages usually bear “cool” aspects due to higher
proportion of deciduous arboreal elements.
For the purpose of the presented paleovegetation maps
several formations have been distinguished and characterized
mostly based on the proportion of broad-leaved deciduous,
broad-leaved evergreen, sclerophyllous and
legume-type components of zonal woody angiosperms
(Kovar-Eder et al. submitted).
Zonal formations
1. (Warm-) temperate Broad-leaved Deciduous Forest
with very low proportion of evergreen woody elements
(vegetation unit 1) includes more than 80 % of zonal deciduous
woody elements of angiosperms, such as Parrotia,
Zelkova, Ostrya, Acer angustilobum etc.
2. Warm-temperate Mixed-Mesophytic Forest (vegetation
unit 2) includes less than 80 % deciduous woody elements of
zonal angiosperms, less than 30 % evergreen broad-leaved
woody taxa of zonal angiosperms and less than 20 % sclerophyllous
and legume type elements, regular admixture of
Tetraclinis salicornioides and other thermophilous elements,
less than 30 % of zonal herbs of zonal angiosperms.
3. Subtropical Broad-leaved Evergreen Forests including
the “Younger Mastixioid Floras” sensu Mai (1964)
(vegetation unit 3) includes equal or more than 30 %
broad-leaved evergreen and thermophilous elements, represented
mainly by Lauraceae, Theaceae, Mastixiaceae,
Symplocaceae, Sapotaceae, Engelhardia, and evergreen
Fagaceae (represented in pollen spectra by morpho-species
Castaneoideoipollenites pusillus, Quercoidites henrici,
Quercoidites microhenrici – types) and less than 25 %
of zonal herbs among zonal angiosperms.
4. Subtropical Sub-humid Sclerophyllous Forest (vegetation
unit 4) includes more than 20 % sclerophyllous taxa
(Quercus mediterranea, Quercus drymeja) and legume-type
microphyllous woody elements of zonal angiosperms.
Intrazonal formations
5. Swamp forest and coal-forming mire (not expressed
by patterns on maps, vegetation unit 7) is dominated by
coal-forming woody and herbaceous elements (e.g. Glyptostrobus
and other Taxodiaceae, Byttneriophyllum, Nyssa,
Myrica, Calamus, Spirematospermum, etc.).
6. Marsh and aquatic vegetation (not expressed by patterns
on maps, vegetation unit 8) is dominated by aquatic
herbs and helophytes (Cyperaceae, Typha, Potamogeton,
Stratiotes, etc.).
7. Deciduous riparian forest (not expressed by patterns
on maps, vegetation unit 9) is dominated by woody elements
of moist substrates (Taxodium, Alnus, Salix, Populus,
Fraxinus, Acer tricuspidatum, etc.).
Extrazonal formations
8. Mountain conifer-rich forest is dominated (mostly in
pollen records, vegetation unit 10) by Pinaceae (including
Cedrus, Tsuga, Picea, Cathaya, etc.).
297MIOCENE EVOLUTION OF LANDSCAPE AND VEGETATION IN THE CENTRAL PARATETHYS
Fig. 1. Geological maps: A – Alpine-Carpathian-Pannonian region, B – Position of the ALCAPA and Tiszia Dacia microplates.
The network of localities is quite loose and does not
sufficiently cover the study area. Thus the gaps between
them have been only tentatively extrapolated according
to the reconstructed relief. The construction of the maps
proceeded in two steps. First, circles of reference sites have
been placed on palinspastic maps that include reconstructed
sea depths and orography. The sites received name abbreviations
(e.g. MA for Mataschen.), which are included
in the explanations of the figures and in the review of the
sites (available in the digital form on request). Different
colours of circles have been used to designate the vegetation
formations. The presence of extrazonal mountain conifers
in the palynospectra has been marked as a bluegreen
rim, all kinds of intrazonal vegetation as the brown
centre or brown full circle. Colours of zonal vegetation
have been divided as follows: light green for the Deciduous
Broad-leaved Forest (unit 1), green for the Mixed Mesophytic
Forest (unit 2), dark green for mostly evergreen
forests (unit 3), and orange for sub-humid, partly sclerophyllous
forest types (unit 4).
In the next step, we used various raster patterns (as specified
in the explanations of the maps) to depict probable
298 KVAČEK et al.
vegetation formations between and around the reference
sites. The intrazonal vegetation was omitted because of its
limited extent compared with the scale of the maps. The
approximate position of extrazonal conifer belts has been
placed according to the nearby palynodata and orography.
Early Miocene – model of the Karpatian landscape
and vegetation of the Central Paratethys
During Early Miocene, the Central Paratethys was situated
at least 200—300 km to the south of its present position.
The sea, located along the southern slopes of the
European platform, covered the southwestward dipping
subduction zone in front of the developing Alpine-Carpathian
orogene. The accretion wedge, build up by external
Alpine and Carpathian Flysch Belt units, was marked
in that time only with islands, forming an archipelago
along the platform margin. The uplifted islands, composed
of units of the Outer Carpathian nappe pile, were surrounded
by residual flysch troughs with turbidite deposition.
Toward the south, the epicontinental sea stretched
over a large part of the internal zones of the Alpine and
Carpathian mountain chains, included the northern lithosphere
fragment of ALCAPA and the southern fragment of
Tisza-Dacia (Csontos et al. 1992; Csontos 1995). The
lithospheric fragments (or microplates) moved towards the
subduction zone separately, their amalgamation set at first
since the Middle Miocene. The southern boundary of the
Central Paratethys Sea was represented by the Dinaride
mountain chain, dividing it from the Mediterranean Sea.
In late Early Miocene, the subduction gradually converted
to a collision from the west to the east. The weight of the
overriding Carpathian orogene front and deep subsurface
load of the submerging plate led to development of a flexure
on the platform margin. The foredeep basin developed
along the whole front of the Outer Carpathian accretion
wedge. The evolution of the accretion wedge was associated
with compression, controlling folding and thrusting of
the Flysch Belt nappe piles (Kováč et al. 1998).
The internal units of the Carpathians, belonging to the
ALCAPA and Tisza-Dacia microplates, started to collapse
due to stretching in consequence of the subduction pull
(Royden 1993a,b), as well as due to asthenospheric mantle
upheaval in the western part of the back-arc region. The
extension led to initial rifting of the Pannonian basin system
(Horváth 1993). By basin opening, besides normal
and low angle faults, strike slip faults also played an important
role (Vass et al. 1988, 1993; Tari et al. 1992;
Fodor 1995; Kováč et al. 1998; Konečný et al. 2002). In
the west a sinistral shear dividing the Alpine and Carpathian
orogenes opened the Vienna Basin, in the east a
dextral shear along the external and internal Carpathians
boundary (Pieniny Klippen Belt) opened the Transcarpathian
Basin towards the northern part of the East Slovak
Basin. Extension in the western part of the back-arc region
led to beginning of the formation of the Danube Basin, associated
with structural unroofing of the deepest AlpineCarpathian
structural units.
During the Karpatian, a new marine connection opened
between the Central Paratethys and the Mediterranean Sea.
This connection is supposed through the trans-Dinaride corridor
situated in the area of Slovenia and northern Croatia
(Rögl 1998). Apart from tectonics, the global sea-level rise
during the late Burdigalian had an important role in the development
of this seaway (TB 2.2 cycle, sensu Haq et al.
1988; Hardenbol et al. 1998; Kováč et al. 2001). The sea
transgression, with new elements of marine fauna and flora,
flooded the present territory of the Drava and Sava Basins
(Pavelić 2001), from where the sea penetrated into the
Mura, Zala and Styrian Basins. The NE oriented flooding
then followed the way between the northern margin of the
Mecsek Mts and the southern margin of the Transdanubian
Range reaching the North Hungarian—South Slovak sedimentary
area. Northwards, the sea spread to the Bánovská
kotlina Depression, the Vienna Basin, the Váh river valley
and the East Slovak Basin (Kováč et al. 1993). The Karpatian
sea covered especially the western part of the Carpathian
Foredeep, in the east the sea extended especially
over the area of the present Outer Carpathian units of accretionary
wedge with wide marine connections into the East
Slovak Basin (Rudinec 1989, 1990; Kováč et al. 1995).
The DEM paleogeographical model of the Central
Paratethys during the Karpatian (Fig. 2) documents the
beginning of the Carpathian orogene uplift. The ratio between
continental and marine environments (ratio of land
and surface covered by the sea) can be very roughly interpreted
due to the enormous erosion of the Early Miocene
sediments at the begin of the Middle Miocene (Kováč et
al. 2003). The erosion is documented by a total absence of
marginal facies particularly in northern areas of the Central
Western Carpathians and by very sporadic findings of
the Karpatian sediments in the Outer Carpathians, folded
together with the Flysch Belt deposits (Cieszkowski 1992;
Oszczypko 2003). The results of study of fluid inclusions
also confirm erosion of 2 to 5 km thick pile of deposits
(Hurai et al. 2002), as well as an important angular unconformity
between the Karpatian and Badenian strata at
many places in the Pannonian basin system.
The Carpathian paleo-relief was probably low at this
time. In many places, the pre-Tertiary basement units were
covered by Paleogene and Early Miocene sediments,
much larger in extent than those preserved today in the
Paleogene and Neogene basins. In the Western Carpathians,
for example, a continuous sedimentary area covering
the territory from the Bánovská kotlina Depression
to the Vienna Basin has been recorded (Kováč et al. 1993),
without indication of an uplift of the Považský Inovec
core mountain (Kováč et al. 1994, 1997).
The paleo-river net started to develop in areas with
higher relief. The Eastern Alps belonged to such places,
where rivers fed deltas in the Alpine Foredeep and the
southern part of the Vienna Basin (Aderklaa Formation),
and later in the uplifted parts of both the Central Western
Carpathians and the Outer Carpathians in the western segment
of the Carpathian collision zone (Kováč 2000).
Considerations about prevailing low paleo-relief are
also supported by the paleobotanical study, which docu-
299MIOCENE EVOLUTION OF LANDSCAPE AND VEGETATION IN THE CENTRAL PARATETHYS
Fig. 2. DEM of the Central Paratethys: landscape & vegetation cover during the Karpatian. Abbreviations of the localities with numbers of
vegetation units: BE – (3, 8) Bełchatów, KRAM-P 211/214; BR – (3, 9) Berzdorf; DD – (3) Dolní Dunajovice, Slup, Hevlín; DE – (3, 7)
Dežerice; GB – (3, 10) Core Gbely 139, depth 650—660 m; HA – (9) Haiden; HI – (3, 9) Core Hidas 53, 1071.0—763.3 m; HR – (3, 7)
Hrádek/N., Kristina Mine, Turów; KA – (3) Kamenný Újezd, Olešník, Hluboká; KL – (3, 10) Kłodnica area, Biała and Twardawa cores;
KO – (4, 9, 10) Core Komló 120; KZ – (3, 9, 10) Komló-Zobák puszta; LA – (3) Laa/Thaya; LE – (3, 10) Leánykö; LI – (3, 9)
Lintsching; LM – (1, 7, 8) Lipnica Mała; MA – (4) Magyaregregy; MO – (3, 7, 8, 10) Modrý Kameň, Stredné Plachtince, Ďurkovce,
Dolné Príbelce; MY – (4) Mydlovary; NO – (3, 7, 9) Core Nosislav 3, 368—345 m; NS – (2, 7, 8) Nowy Sącz; PA – (4, 9) Parschlug;
PE – (9) Core Pécsvárád 44; PI – (3, 8, 9) Core Piliny 8; PU – (3, 8, 10) Core Püspäkhatran 4; TE – (3, 9) Core Tekeres 1; TR – (2, 8, 9)
Teiritzberg; VA – (3, 9, 10) Core Várpalota 133; ZA – (7) Zangtal near Voitsberg; ZD – (3, 9, 10) Cores Ždánice 67, depth 765—858 m,
Ždánice 68, depth 700—820 m; ZE – (3, 9, 10) Core Zengővárkony 59; ZO – (9, 10) Core Zohor 1, depth 1495—1500 m.
300 KVAČEK et al.
ments an altitudinally not much differentiated character of
vegetation in the whole Carpathian-Pannonian region during
the Karpatian. Similarly, no obvious latitudinal and
longitudinal changes were observed there. Some zonation
marks, indicated by the diferentiation of the paleovegetation
cover, are visible in the lowlands and margins of the
marine sedimentary area.
Karpatian vegetation (Fig. 2) is documented by reference
sites, of which only some have been dated by marine
fauna. The age of the others is indicated by the mammalian
MN5 Zone and thus may overlap with the Early Badenian.
In other cases the boundary to the Ottnangian may
also be uncertain (MN4 Zone).
Three zonal forest formations were spread over the Central
Paratethys region during the Karpatian. Subtropical
broad-leaved forests with high to medium proportion of
evergreen elements were spread in the western part and
continued into the Boreal Province westwards in the form
of the typical Younger Mastixioid Floras sensu Mai
(1964). The corresponding phytostratigraphic unit of the
Boreal Province has been called the Floral Assemblage
(“Florenkomplex”) Františkovy Lázně – Kleinleipisch
(Mai 1995, 2001; Czaja 2003), but direct dating to the
lower part of the MN5 Zone is available only for Františkovy
Lázně in the Cheb Basin outside the Central Paratethys.
Domination of thermophilous elements (Engelhardia, Platycarya,
Sapotaceae, Symplocaceae, evergreen Fagaceae,
such as Trigonobalanopsis, in pollen records expressed by
Castaneoideaepolis pusillus-, Castaneoideaepolis oviformis-,
Tricolporopollenites liblarensis-, Quercoidites henrici-
types, etc.) is apparent for the Karpatian—Early Badenian
time span (Planderová 1990, Zone MF5; Doláková &
Slamková 2003). Only in the central and northern parts
near mountains (e.g. Lipnica Mała, Nowy Sącz), the proportion
of the broad-leaved deciduous elements increased
resulting in the warm-temperate Mixed Mesophytic and
Broad-leaved Deciduous Forest types. Some sites (Mydlovary
– Knobloch & Kvaček 1996; Parschlug – Kovar et
al. 2004; Magyaregregy – Hably 2002) have a sub-humid
character (subtropical forests with high proportion of
sub-humid and sclerophyllous elements). This may indicate
some heterochronity, which we are unable to resolve
from the paleobotanical record and short-time fluctuation
of perhumid and seasonal climate, as proposed by exothermic
vertebrates (Böhme 2003). A distinct East-West gradient
is apparent, when broader parts of Europe are
compared (Kovar-Eder et al. submitted).
Extrazonal mountain zones with conifers were probably
as high as 1500—2000 m a.s.l. and more. The upland forests
were dominated by Pinus, Abies, Cathaya and only at
still higher altitudes with the admixture of Cedrus, Tsuga,
and Picea, as demonstrated in the pollen spectra (Nagy
1992; Doláková & Slamková 2003). Various Pinaceae (Pinus,
Cathaya) and Sciadopitys entered also intrazonal
lowland formations.
Intrazonal coal-forming forests appeared mainly in the inter-Alpine
basins (Leoben-Bruck Basin, Parschlug, Fohnsdorf,
and Mecsek Mts – Hably 2002, p. 92). Besides
Glyptostrobus these thermophilous communities occasionally
included palms, e.g. Calamus-type at Teiritzberg, evergreen
oaks, Nyssa, Myrica, Cyrilla (Zittau Basin), and not
yet clarified enigmatic Rhoipites pseudocingulum (= Rhustype).
Among other mostly intrazonal elements, ferns of
Gleicheniaceae, Schizaeaceae (Lygodium), Osmunda and
Polypodiaceae sensu lato (including, e.g. Pronephrium)
were well represented. Pollen of Avicennia (Korneuburg Basin)
suggests the presence of impoverished mangrove
shrubs in the NW part of the Central Paratethys.
Paleobotanical data are lacking for this time interval from
the eastern part of the studied region because of unfavourable
conditions for the preservation of fossil plants there
(Syabryaj 2003). Assemblages from the upper part of the
Smoliarka Horizon (Rylova et al. 1999) and Rozhok (zone
IV FC sensu Yakubovskaya 1993) in Belarus give evidence
that the subtropical vegetation of the mastixioid type may
have extended north-eastwards from the Central Paratethys.
Middle Miocene – model of the Late Badenian
landscape and vegetation of the Central Paratethys
During the Middle Miocene, the active collision/subduction
in front of the Carpathians shifted eastwards due to
gradual break-down of the submerging slab (Tomek & Hall
1993). In the west, this process led to the termination of collision
between the orogene and the European platform, followed
by gradual uplift of the Outer Carpathian
accretionary wedge and by the migration of foredeep depocentres
from the Western Carpathian foreland towards the
Eastern Carpathians (Jiříček 1979). The sea flooding in the
front of the Carpathian orogene gradually schifted from
west to east. At the end of the Early Badenian, the sea abandoned
the western part of the foredeep (Czech Republic),
marine sedimentary areas extended only in the northern
front of the Western Carpathians and in the front of the
Eastern Carpathians. The foredeep at the edge of the Western
and Eastern Carpathians reached its maximum extent
during Late Badenian—Early Sarmatian time, when 2500 m
of sediments were deposited (Meulenkamp et al. 1996). The
Sarmatian compression associated with uplift of the Outer
Western Carpathians draw the sea away from the northern
part of the Carpathian Foredeep for ever (Kováč et al. 1998).
A connection between the Carpathian Foredeep and the
Pannonian basin system remained preserved only in the
Eastern Carpathian region (Kováč 2000; Kováč et al. 1998).
The Middle Miocene development of the back-arc basin
region was controlled by two geodynamic factors: in the
western and central parts of the Pannonian basin system it
was upheaval of asthenosphere mantle masses, in the eastern
part it was stretching of an overriding plate induced by
subduction pull in front of the Eastern Carpathians. In the
western part of the back-arc basin, subsidence of the Vienna
and Danube Basins caused depocentres above the thinned
crust and lithosphere associated with volcanic activity in
the hinterland of the Central Western Carpathians (Wernike
1985; Nemčok & Lexa 1990; Tari et al. 1992; Konečný et
al. 2002). In the east, orogene that was parallel to the backarc
basin depocentres opened in the area of the Transcar-
301MIOCENE EVOLUTION OF LANDSCAPE AND VEGETATION IN THE CENTRAL PARATETHYS
pathian and Transylvanian Basins. Mighty acid and later
also “island arc” type volcanic activity appeared in a belt
along the eastern border of the back-arc region in the hinterland
of the Eastern Carpathians (Konečný et al. 2002).
Synrift subsidence of the back-arc basin in the Pannonian
basin system was associated with mighty acid and calc-alkaline
volcanic activity (Kováč 2000; Konečný et al.
2002). Individual depocentres formed in the extensional
tectonic regime. The main types were grabens and half-grabens
associated with normal and low angle faults, although
some pull apart basins opened along active strike-slip faults
(Vass et al. 1988, 1993; Tari et al. 1992; Fodor 1995; Kováč
et al. 1998; Konečný et al. 2002; Kováč 2000).
The Badenian marine conection of the Central Paratethys
with the Mediterranean Sea is supposed through a
trans-Dinaride corridor (Rögl 1998), at a similar place as
during the Karpatian. The Early Badenian sea-level rise,
which can be correlated with the early Langhian global
sea-level change (TB 2.3 cycle sensu Haq et al. 1988;
Hardenbol et al. 1998), is documented only in the SW part
of the Pannonian basin system – from the Styrian Basin
(Rögl et al. 2002). In the northern parts of the back-arc region
wide-ranging erosion of Early Miocene deposits is
observed. In depressions (future basin depocentres), fan
deltas or terrestrial red coloured sediments were deposited
during this time.
The Early Badenian tectonically-controlled transgression,
followed by rapid subsidence (deep sedimentary environment),
started about 15 Ma. The basins were filled by
clastics transported by rivers running from uplifted areas
of the Eastern Alps and the Western Carpathians. After sealevel
fall (documented by erosion of the Early Badenian
carbonate platforms in the SW part of back-arc basin – Vienna
and Styrian Basins), the “Middle Badenian” transgression,
which can be correlated with the global sea-level
change in late Langhian (TB 2.4 cycle sensu Haq et al.
1988; Hardenbol et al. 1998), took place. The Central
Paratethys sea reached the present extent in the intra-Carpathian
Neogene basins, except the uplifted North Hungarian—South
Slovak sedimentary area. Since that time, a
gradual filling up of the Pannonian basin system by deltas
has been observed, leading to shallowing of sedimentary
environment and development of isolated depocentres. Basins
situated in the north and east suffered from isolation.
A salinity crisis took place in the Carpathian Foredeep as
well as in the Transcarpathian and Transylvanian Basins.
The Late Badenian transgression, which can be correlated
with the global sea-level change at the Langhian/Serravalian
boundary (TB 2.5 cycle, sensu Haq et al. 1988; Hardenbol
et al. 1998; 13.65 Ma.) represents the last full marine
flood of the Central Paratethys. Since the end of the Late
Badenian (12.7 Ma), an isolation of epicontinental sea in
the Intra-Carpathian region can be documented and a direct
connection of the Central Paratehtys with the Mediterranean
is not expected. The Sarmatian flood, from the Eastern
Paratethys region, can be correlated with the late Serravalian
global sea-level change (TB 2.6 cycle sensu Haq et al.
1988; Hardenbol et al. 1998). The sedimentary environment
was dominantly shallow marine with decreased salinity.
The DEM model of the Central Paratethys during the
Late Badenian (Fig. 3) documents uplift of the Western
Carpathians, including accretionary wedge in front of the
orogene and broad marine flood in the back-arc basin region.
The intra-Carpathian region gained the characteristic
features of an archipelago sea, with many small islands
surrounded by shallow epicontinental sea. The paleogeography
was still strongly influenced by tectonic processes
(Styrian phase), as it is well marked by rapid
changes of subsiding depocentres (sea bays, small basins)
and position of the coastal line.
The paleo-relief of the Carpathians significantly changed
during the Middle Miocene, due to strong tectonic influence.
The belt of the Outer Carpathians, bordered by internal
parts of the orogene, started to be uplifted. The land
surface was also strongly differentiated by volcanic activity,
some stratovolcanoes reached heights of 2000 to
3000 m a.s.l. The river net transporting eroded clastic material
headed mostly towards the back-arc area.
Fair-sized altitudinal differences between lowlands and
mountains are documented by paleobotanical studies.
Mixed pollen spectra with mountain and lowland vegetation
taxa indicate only seemingly the decrease of thermophilous
taxa and the increase of more dominant temperate
taxa (Sitár & Kováčová-Slamková 1999; Slamková 2004).
The Badenian vegetation in the Central Paratethys can
generally be characterized as thermophilous without dominance
of typical boreal plant elements. Hence, we cannot
document sufficiently a gradual cooling of climate indicated
in the literature (Böhme 2003) that influenced the
European flora from the Late Badenian onwards.
Late Badenian vegetation (Fig. 3) has been documented
from sites more often dated by marine fauna (due to an
extensive marine transgression), but also from some others
that are dated by mammals to Zone MN6 or by regional
correlation. The differentiation of the levels within the
Badenian has not always been accomplished and such
sites are exceptionally included into the map. Some sites
overlap with the lowermost Sarmatian, which is floristically
hardly distinguishable (see Syabryaj & Stuchlik 2004;
merged into a single “Florenkomplex” Stare Gliwice-Unterwohlbach
by Mai 1995).
The southernmost sites in Romania, Serbia and Hungary
differ from the remaining ones by thermophilous, partly
sub-humid aspects under subtropical climatic conditions,
which continued from previous times. A general cooling
trend appears in other sites by an increasing role of deciduous
elements, while thermophilous plants withdrew stepwise
southwards and only a part survived, mainly
Tetraclinis, Amentotaxus, Magnolia, Lauraceae, Engelhardia
and others. This is in the contrast to the preceding
“Wieliczien” thermophilous humid mastixioid assemblage
even in the Polish part of the Carpathian Foredeep (Łancucka-Środoniowa
& Zastawniak 1997). From the differentiation
of vegetation in the Late Badenian it is obvious
that the climatic gradient between the southern and northern
parts of the Central Paratethys increased at that time,
partly due to altitudinal differenciation, as noted above.
A noteworthy forest-forming tree was Fagus accompa-
302 KVAČEK et al.
Fig. 3. DEM of the Central Paratethys: landscape & vegetation cover during the Late Badenian. Abbreviations of the localities and
numbers of vegetation units: AL – (2) Core Alsóvadász 1; BU – (1, 9) Burkalo; CI – (3) Ciocadia; DNV – (2, 7, 10) Devínska Nová
Ves – brickkiln; ET – (4) Eger-Tihamér; GK – (2, 8, 9, 10) Gdów area, core Kłaj 1, depth 30—405 m; HI – (2, 10) Core Hidas 53;
HN – (2, 7, 9) Handlová-Nováky; KO – (1, 9) Kolisky; KS – (1, 9) Kosov; MS – (1, 7) Myshin; NO – (2, 9) Nográdszakál, Páris
valley; PI – (3) Pirlage; PS – (3) Pistynka; SE – (4) Selishte; TE – (2, 10) Core Tengelic 2; VE – (1, 9) Verbovets; ZA – (2, 9) Zalescy.
303MIOCENE EVOLUTION OF LANDSCAPE AND VEGETATION IN THE CENTRAL PARATETHYS
nied by other deciduous Fagaceae. Acer was diversified
in several species, among which Acer aegopodifolium
(syn. A. quercifolium – see Ströbitzer-Hermann 2002;
Walther & Zastawniak 2005) appears for the first time in
the Central Paratethys (Kovar-Eder et al. 1994), together
with Ginkgo, Eucommia both arriving from Asia via the
Turgay migration route. Over most of the Central Paratethys,
warm-temperate Mixed Mesophytic Forest thrived,
only in the Western Carpathians and the Transcarpathian
Ukraine some sites already acquired the character of
Broad-leaved Deciduous Forest, probably due to influence
of mountains or due to cooling effect of intensive volcanic
activity (Navrotskaja et al. 1991; Syabryaj 1992). The
proportion of herbs was generally low, not indicating lowland
open vegetation (Syabryaj & Stuchlik 2004).
Extrazonal mountain forests are well discernible in pollen
assemblages, as at Devinská Nová Ves (Sitár &
Kováčová-Slamková 1999), where the coniferous belt reflected
by the dominant pollen of Pinus includes additional
high-mountain elements, such as Cedrus and
Tsuga. Its lower boundary may have decreased towards
1200 m a.s.l. However, this guess is only inferred from the
analogous Recent situation in the Colchis area (Stuchlik
& Kvavadze 1987; Klotz 1990).
In the intrazonal Glyptostrobus peat-forming forests,
Byttneriophyllum and Alnus constituted a basic community,
which became wide spread later in the Neogene.
Among riparian elements, Platanus leucophylla is occasionally
present. Lignite-forming communities are typically
developed in intra-montane depressions (e.g. Handlová
and Nováky) and they are also common in the lowlands
outside the Paratethys area in Poland and Germany (Lusatia
seam 1). The corresponding phytostratigraphical level
in the Boreal Province is probably represented by the Floral
Assemblage Schipkau-Konin sensu Mai (2001), but
this correlation is opposed by Krutzsch (2000.) The paleofloristic
differentiation around the Late Badenian and Early
Sarmatian boundary has been discussed with little
success to give clear-cut differences based on plant
megafossils (Němejc 1951, 1967; Shvareva 1965; Sitár
1967, 1982). It is still uncertain, which Early Miocene elements
did not enter the Sarmatian flora, where broadleaved
deciduous trees predominate. In her pollen
assemblages, Planderová (1990) created a transitional
Zone MF7 for this type of flora in the Slovak Neogene. Its
pollen spectra include a very low proportion of or no thermophilous
Symplocaceae, Sapotaceae and Cyrillaceae.
Late Miocene – model of the Middle Pannonian
landscape and vegetation of the Central Paratethys
During the Late Miocene, the area of the Central Paratethys
gained paleogeographical features similar to the
present situation in the Carpathian-Pannonian region. The
main difference was represented by flooding of the intraCarpathian
region and the foredeep depocentres, which
were restricted to the southeastern foreland of the Eastern
Carpathians in that time (Jiříček 1979; Meulenkamp et al.
1996). Thus a connection of the brackish Eastern Paratethys
with the Lake Pannon originated (Magyar et al. 1999).
The Late Miocene geodynamic development of the Carpathians
can be characterized by termination of collision
between the Western Carpathian and the European platform
and reinforced collision connected with subduction
in front of the Eastern Carpathians. This process was followed
not only by the uplift of the accretionary wedge
loop, but also by the uplift of the whole Carpathian orogene
mountain chain.
Pull of the active subduction in front of the Eastern Carpathians
southern edge led to the “second” rifting phase
in the back-arc basin at the begining of the Pannonian,
followed by thermal post-rift subsidence (Lankreijer et al.
1995; Kováč 2000; Konečný et al. 2002). Among the Late
Miocene basins of the intra-Carpathian domain formed in
an extensional regime, flexural basins without important
fault activity prevailed, although in the Early Pannonian
normal and strike slip faults allowed development of small
pull-apart basins (Vass et al. 1988, 1993; Tari et al. 1992;
Fodor 1995; Kováč et al. 1998; Kováč 2000; Konečný et
al. 2002). The Lake Pannon – the Pannonian basin system
was gradually filled up by deltaic deposits, generally
from the northwest toward the southeast. The sedimentary
environment gradually changed from a brackish deepwater
to a shallow water – lake environment due to isolation
from the Mediterranean and Eastern Paratethys (Magyar et
al. 1999). At the northern margin of the Lake Pannon,
marshes, swamps and deltaic systems spread in an everlarger
extent. Due to the retreat of the coastal line the
lacustrine environment changed generally into alluvial
also in the Late Pannonian. Later, in the Pliocene a broad
area of lowlads appeared in the hinterland of the Carpathian
chain. Scattered mountains (e.g. the Transdanubian
Range Mts, Bükk Mts, Apuseni Mts) and basalt
volcanoes formed higher morphological elevations.
At the end of the Late Miocene, tectonic inversion of
the basin system (Horváth 1993, 1995; Horváth & Cloetingh
1996) led to the retreat of the aquatic sedimentary
environment in the whole intra-Carpathian area, exept the
central and southeastern regions. Uplift of the Eastern
Alps, the Western and Eastern Carpathians was associated
with angular unconformity between the Middle and Late
Miocene (or younger) strata.
The architecture of the Late Miocene fill of the Pannonian
basin system was distinctly influenced by paleogeographical
changes. From the sequence stratigraphy and
depositional systems point of view, the succession of the
Late Miocene sediments can be characterized at first by a
dominant portion of proximal deltaic deposits (A—C zones,
sensu Papp 1951), which pass upward into distal deltaic
to basinal fine-grained clay—silt—sandy facies (D, E zones).
Fluvio-lacustrine sediments with coal seams (F—H zone)
formed the terminal part of the Pannonian. The overlying
Pliocene strata were deposited mostly in an alluvial sedimentary
environment. The Pannonian cyclicity can be correlated
by global changes Tor-1 cycle (11.6—9.3 Ma, sensu
Hardenbol et al. 1998) and Tor-2 cycle (9.3—7.2 Ma, sensu
Hardenbol et al. 1998).
304 KVAČEK et al.
The DEM model of the Central Paratethys during the
Middle Pannonian (Fig. 4) documents the extinction of
the sedimentary area of the Carpathian Foredeep, except
its southeastern part and uplift of the Carpathian mountain
chain. At that time, a maximum extent of brackish seawaters
covered the intra-Carpathian domain.
The expected paleo-relief of the Carpathian orogenic
chain in the Late Miocene began to match the present-day
situation, characterized by the presence of both mountains
and lowland areas. A difference can be seen in the more elevated
belt of the Outer Carpathian accretionary wedge,
which formed a natural barrier between flat territories of
the European platform and the ever-subsiding Pannonian
basin system covered by the Lake Pannon. During the
Pannonian and Pliocene, basalt volcanic activity also participated
in paleo-orography of the back-arc region
(Kováč et al. 1998; Kováč 2000).
The Middle Pannonian vegetation (Fig. 4) corresponds
to the warm temperate climatic zone with evidence of sporadically
present termophilous and evergreen taxa. A higher
percentual proportion of non-arboreal pollen (10 to 14 %)
indicates local vegetation of partly open woodland (i.e.
woods with open canopy). An increase of halophytic taxa
documents the presence of coastal lagoons and marshlands
during the lowstand of the brackish sea. Swamp vegetation,
which grew directly on swamp substrates, is characterized
mainly by noteworthy Taxodiaceae trees. They are often
present in the association with Myricaceae and subordinary
Nyssaceae. The riparian forest elements subdominantly occurred
with Alnus and Ulmus. The extrazonal vegetation of
the mountain areas with Picea, Tsuga, Abies, Cedrus is well
represented in the pollen spectra.
During the Late Pannonian, the Western Carpathian paleogeography
started to change. The Lake Pannon withdrew
southwards, the nothern margin of the back-arc basin was
slightly uplifted with the progradation of deltaic and alluvial
facies, especially in lowlands. These areas were often covered
by hygrophilous plants: Myrica, Salix, Ulmus, Alnus. Herbs
were represented by Chenopodiaceae, Asteraceae, Ericaceae,
Poaceae and Artemisia. Unevenly high moutain relief of the
uplifted mountain chains created ideal conditions for the
mixed mesophytic forests with Carya, Quercus, Craigia,
Carpinus, Fagus, Picea, Abies, Tsuga and Pinus.
The reference sites considered on the map have been assigned
to the Pannonian zones C—E sensu Papp 1951,
namely the time slice before the major spreading of the
lignite facies over the Pannonian Basin (zone F). The dating
is based mostly on molluscs or regional correlation,
rarely on mammals (MN9 Zone). Additional sites of Early
Pannonian age (zones A—B) are shown in brackets on the
map. They indicate the previous vegetation type at the
Sarmatian/Pannonian boundary. Most leaf assemblages of
Pannonian age are at least partly intrazonal and it is difficult
to obtain a true picture of zonal vegetation from them.
Most spore/pollen assemblages, mainly from the Hungarian
Pannonian, have not been revised and the stratigraphic
position of pollen samples remains partly uncertain.
At the very beginning of the Pannonian, subtropical conditions
returned to some parts of the Central Paratethys. The
thermophilous vegetation from southern Austria (Mataschen,
Styrian Basin, Pannonian B – Kovar-Eder 2004) can also be
found to the north-western periphery outside the Paratethys
(Gozdnica) and may correspond to the Floral Assemblage
(“Florenkomplex”) Düren sensu Mai (1995) in western Europe
with the latest occurrences of mastixioid plants. However,
the dating of Gozdnica is still under dispute (see Dyjor et
al. 1992, 1998; Mai, personal communication).
Later in the Pannonian, thermophilous evergreen, partly
sub-humid vegetation remained in the south and south-eastern
parts (Serbia, southern Hungary, the Borod Basin in Romania).
In the Middle Pannonian, broad-leaved deciduous
and Mixed Mesophytic warm-temperate to temperate forests
with a low proportion of evergreen elements generally
became widespread over the Central Paratethys (Styrian Basin,
Molasse Zone, Vienna Basin). Characteristic elements
in these communities included Fagus haidingeri—pliocenica
complex, Quercus (?Castanea) kubinyii, Quercus
pseudocastanea—pseudorobur complex, Carpinus sp. div.,
Betula, Acer integrilobum, Acer vindobonense and Acer
subcampestre (syn. Acer jurenakyi) (Kovar-Eder 1988;
Ströbitzer-Hermann 2002; Ströbitzer-Hermann & KovarEder
2003). Cooling trends can be traced in spore/pollen
spectra from the cores in the Pannonian Basin (Nagy &
Planderová 1985; Nagy 1992). At some sites (Alsóvadász,
Suchohrad) somewhat higher frequencies of Chenopodiaceae
and Artemisia (max. 15 %) may indicate patches of
herbaceous vegetation on marshes within broad-leaved
deciduous forests. However, high mean annual precipitation
(Bernor et al. 2003) prevented the development in
this aera of open woodland and grassland vegetation like
that suggested for south-eastern to southern Ukraine outside
the Central Paratethys (Syabryaj 1999, 2003). Temperate
purely deciduous broad-leaved forests were also
spread during the Middle Pannonian in the Western Carpathians
(e.g. sites at Nové Ustie in the Orava Basin and
Martin in the Turiec Basin). In most cases these assemblages
include a considerable proportion of intrazonal elements,
both woody and aquatic (Trapa). Characteristic
riparian woody elements were Salicaceae, Platanus leucophylla,
Alnus ducalis, Alnus cecropiifolia, and partly intrazonal
Quercus gigas and Pterocarya paradisiaca. The
dominant peat-forming community during the whole Pannonian
consisted mostly of the Glyptostrobus-Byttneriophyllum-Alnus
swamp forest. This swampy coal-forming
vegetation reached its widest distribution over the Pannonian
Basin during the Late Pannonian, as the large extension
of the lignite facies in Hungary, southern Moravia,
Slovakia and Serbia corroborates (Knobloch 1969; Givulescu
1992; Pantić & Dulić 1993; Hably 2003).
Extrazonal montane conifers of all kinds (including
Tsuga, Cedrus, Picea, Abies, Keteleeria) were found in
pollen spectra throughout the Pannonian Basin as regular
accessories. In the intra-montane basins macrofossils of
these conifers were also formed (Nove Ustie, Martin).
Therefore we may expect high mountain conifer belts
reaching over 1500 m a.s.l. on the Carpathian and Alpine
ridges and descending to medium altitudes, mixed with
deciduous broad-leaved elements.
305MIOCENE EVOLUTION OF LANDSCAPE AND VEGETATION IN THE CENTRAL PARATETHYS
Fig. 4. DEM of the Central Paratethys: landscape & vegetation cover during the Middle Pannonian. Abbreviations of the localities (in
brackets earliest Pannonian, not considered for raster pattern) and numbers of vegetation units: AL – (1, 10) Core Alsóvadász 1, depth
155.8—240 m, Cszerehát environment; BE – (1, 8, 9, 10) Belchatow, upper level (section VI.1, KRAM-P 17, Stawek-1A); BO – (9, 10)
Bobrov, core V 6; DE – (3) Delureni; DU – (4, 9) Dubona I; EB – (9) Ebersbrunn; (GO) – (3, 8, 9, 10) Gozdnica; HI – (2, 8, 9, 10)
Core Hidas 53, depth 298—367 m; (HO) – (8, 9) Höllgraben; KO – (9) Kogelwald, core KO 4; (KU) – (7, 8) Kunovice, cores KU 1,
depth 42-46 m, KU 2, depth 109—174 m; LA – (2, 9) Laaerberg; (MA) – (3, 8) Mataschen near Fehring; ME – (1-2, 9, 10) Core Megyaszó
1, depth 52—206 m; MI – (2, 8) Mistřín, DV 4 Mine; MR – (1, 8, 9) Martin, Turiec Basin; MU – (7, 9) Münzengraben, core MÜ 21;
(NE) – (3) Neuhaus/Klausenbach; NI – (1, 8, 10) Nitra environs, Vozokany, core N-7, Rohoznica, core N-8, Mechenice, core B-23, Pohranice,
core B-25; NU – (8, 9) Nové Ustie, Orava Basin; OR – (7, 8) Ořechov, Polešovice, cores UH 18, depth 11.3 m, UH 19, depth
14.1—29.4 m; (PA) – (2, 8, 9) Paldau; PO – (7, 9) Pöllau, core PÖ 2; RE – (3, 8, 9) Reith near Unterstorcha; RU – (2, 7, 9) Rudabánya;
SK – (3, 9) Sremska Kamenica; SO – (1, 8, 9) Sośnica; (SU) – (2, 9, 10) Core Suchohrad 32, depth 625—638 m; TA – (1, 9, 10) Tata,
Core TVG 26 depth 7—39m; TO – (1, 9, 10) Core Tököl 1, depth 688.5—730 m; (VC) – (3, 4) Valea Crisului; VO – (2, 8, 9) Vösendorf;
WO – (8, 9) Wörth near Kirchberg/Raab.
306 KVAČEK et al.
Paleoclimatic trends
Palaeoclimatic proxies are available for many sites considered
in the presented maps (see Mai 1995). Most of
them have been derived from “intuitive” comparisons
with extant vegetation, while the objective co-existence
and leaf physiognomical (CLAMP) methodologies have
not been largely applied so far.
Broad-leaved evergreen forests, including the “Late Mastixioid
Floras”, were usually compared with similar forests
in monsoon East Asia. In general, the climate corresponding
to this type of vegetation was humid to per-humid (annual
sum of precipitation about 1000 to 3000 mm), with heavy
rains prevailing in the summer, but without any month with
a humidity deficit. The temperature of the coldest month
varied between 4—10 ºC and absolute minima rarely
reached under zero. The Mean Annual Temperature (MAT),
according to various estimates (e.g. Mai 1995) varied in
larger spans, according to the percentage of evergreen taxa,
from 13 to almost 20 ºC. The climate must have been quite
equable, with a range of temperature less than 25 ºC. This is
a prevailing condition in the Karpatian to Lower Badenian
stages over the Central Paratethys. Only in some parts, high
summer mean temperatures over 20 ºC and also dry substrate
caused a relative humidity deficit and local expansions of
sub-humid sclerophyllous (microphyllous) evergreen forests.
This type of forest has nothing to do with the etesian (Mediterranean-type)
climate and was probably more similar to
the microphyllous montane forests in drier parts of the Himalayas
today.
The cooling trends expressed by the decline of MAT in
the Late Badenian over northern and eastern parts of the
Central Paratethys caused changes in the forests. Evergreen
elements mostly withdrew southwards and only less
frost-sensitive trees remained forming the Mixed Mesophytic
Forest with high representation of deciduous taxa.
MAT, under which this type of vegetation optimally
thrives, is variously estimated, depending on what type of
forests is brought for comparison – East Asiatic or North
American (Wolfe 1979). While in the former, the Coldest
Month Mean Temperature (CMMT) is usually quite low
(up to —2 ºC), the latter thrives in warmer, subtropical conditions
(northern Florida – mean annual temperature ca.
20 °C, January mean up to 10 ºC). In general, the following
estimation of the decline in temperature can be expected
for the Central Paratethys from the published data
(e.g. Mai 1995) – MAT 16(—?10) ºC, that is lowering by
about 3 ºC in comparison with the Karpatian, and adequate
lowering of the January mean, which may have been
even stronger, because of decrease of climatic equability.
The sum of Mean Annual Precipitation (MAP) remained
high enough for humid conditions in any case, also thanks
to the colder climate. Such warm temperate conditions
predominated during the Late Badenian and the Middle
Pannonian. In the Pannonian in northern and easterly parts
of the area studied, particularly near the mountains, the
Deciduous Broad-leaved Forests indicate still more severe
deterioration of climatic conditions. These temperate forests
dominated by deciduous oaks and beech and intermixed
with various Pinaceae withstood decrease of
CMMT up to —10 °C, particularly on higher altitudinal
habitats. The impact of such severe winters was certainly
milder due to heavy snows, as is the case today in East
Asia, particularly in Japan. Equally high precipitation
throughout the year also prevented expansion of herbaceous
steppe vegetation, although patches of it can be noticed
in the pollen record within deciduous broad-leaved
forests in the Middle Pannonian. Due to high humidity of
climate, extensive lignite deposits originated over most of
the Paratethys area and its north and west periphery.
Conclusions
After having compiled the presented Miocene geobotanical
maps according to the methodology applied above, the
following conclusions can be drawn for the future research
in other areas and time slices of the Cenozoic.
Contrary to various models of ancient Cenozoic vegetation
that rely on the physiognomy and composition (diversity)
as well as abundance of elements (e.g. Wolfe
1979; Mai 1995), the presented system of vegetation units
is much more simplified. It surely suffers from various deficiencies.
It neglects abundance. But this parameter is, in
our opinion, not objectively derivable from frequencies of
fossils in a given site or core level. Another weak point of
the system employed above is that individual elements
with a broader ecological span can enter more units or
they are transitional and their foliar physiognomy (evergreen
vs. deciduous) cannot be precisely identified (e.g.
Symplocos, Engelhardia). The defined vegetation formations
were certainly not profoundly clear-cut in ancient
landscapes and transitions between them existed. Still the
diversity percentages are most objective characteristics for
a given assemblage and can be verified any time, the assignment
into the system of the defined vegetation formations
as characterized above (see also Kovar-Eder et al.
submitted) is easy and mostly unequivocal.
According to our experience, it is advisable to use paleogeographical
and geobotanical maps of narrow time
slices, because they reveal better consistent patterns of
vegetation and its dynamics in spite of fewer reference localities.
When a longer time interval has been considered,
local differences between the sites sometimes expressed
trends in time rather than climatic gradients in space (see
Fig. 3 for the Early-Middle Pannonian).
Megafossil and spore/pollen plant records were combined,
whenever feasible, to gain better understanding of
taxonomy and ecology of elements composing assemblages.
Of course, great problems exist, concerning how to
transfer spore/pollen diagrams with various enigmatic
plant elements into a formation with known physiognomy.
In the fututre, it would be desirable to unify views on
the taxonomy of those elements of uncertain affinities
both in megafossil and spore/pollen records.
Older data of paleobotanical and palynological research
have not been neglected but revised and transferred to a
common nomenclature, when the documentation (illustra-
307MIOCENE EVOLUTION OF LANDSCAPE AND VEGETATION IN THE CENTRAL PARATETHYS
tions, preparations) was available and re-studied. In many
cases, such assemblages and their elements were wrongly
interpreted, or wrongly assigned to the natural system. Yet
the current progress in knowledge of whole fossil plants is
improving these inconsistencies. Actuopalynological studies
of Recent vegetation (e.g. Stuchlik & Kvavadze 1987;
Kvavadze & Stuchlik 1990, 1993) offered important clues
for converting spore/pollen spectra into various types of
real vegetation. It would be worth attempting to apply for
the same sets of fossil data the new methodology presented
in this account, which is based on proportions of
components and diversity, along with that currently employed
by palynologists, which uses abundance percentages
of elements.
Acknowledgments: We appreciate discussion, data and
exchange of views with our colleagues L. Stuchlik,
Kraków, V. Mosbrugger and A. Bruch, Frankfurt am
Main and I. Magyar, Budapest. L. Stuchlik, L. Hably and
D. Ivanov as the reviewers and P. Bosák as the editor who
suggested useful improvements to the first version of the
text. This research has been sponsored by the European
Science Foundation (Project EEDEN), by the Czech
Grant Agency GAČR (Project No. 205/04/0099), Slovak
Grant Agency VEGA (Projects No. 2/5016/05, 1/2035/05,
1/0080/03), project of the Slovak Research and Development
Support Agency APVV-51-011305 and the Slovak
Ministry of Education (Project No. AV/808/2002).
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The Holocene
2016, Vol. 26(9) 1345­–1354
© The Author(s) 2016
Reprints and permissions:
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DOI: 10.1177/0959683616640051
hol.sagepub.com
Introduction
The period of the so-called Last Glacial Maximum (LGM), which
is part of the Late Pleniglacial (e.g. Magyari et al., 2014a, 2014b),
is defined on the basis of the regression of sea level (Yokoyama
et al., 2000) or the related maximum extent of glacier ice (Clark
et al., 2009; Ehlers et al., 2013), which occurred between 26.5 and
20/19.0 kyr cal. BP. At the same time, the LGM period is generally
considered to be climatically extreme, conditions in which the
hitherto known Palaeolithic settlements retreat from exposed areas
into preferential (more sheltered) refuges. In Germany, for
instance, this period is usually described as the time of resettlement
of the original occupied areas that became re-colonised only
with the onset of the Magdalenian permeating from France (Terberger,
2013; Terberger and Street, 2002). Similarly, the reduction
in the number of significant sites has also been noted in Moravia in
the past (Oliva, 2005; Svoboda et al., 2002; Valoch, 1996, 2010).
The climatic conditions considerably influenced Central,
Western and Northern Europe during the highest phase of the
Weichsel glaciation. The palaeoclimate in the Middle Danube
region during the LGM is reconstructed as mean summer temperature
of about 11°C with mild September (ca. 7°C). Winter
began in November (−5°C), and in January and February, the temperature
dropped down to −11°C to 12°C. The proposed mean
annual precipitation peaked in July and August (3 mm/day), then
it fell to its annual minimum of 1 mm/day in October, staying at
an average 1–2 mm/day through winter (Davies et al., 2003). The
Czech Republic and the adjacent territories were part of a relatively
narrow (about 400 km wide) zone delimited by the front of
the continental glacier in the north and by the Alps in the south
(extent of glaciation, e.g. Ehlers et al., 2013).
The reconstruction of Late Pleniglacial vegetation has traditionally
been based on interpreting pollen data. Many pollen
records of the period of the Last Glacial in Central Europe show a
shift in pollen from dry steppe/shrub tundra to a birch/juniper
woodland followed by a birch/pine forest (e.g. Amann et al.,
2013; Jankovská and Pokorný, 2008; Magyari et al. 2014a, 2014b;
New information augmenting the picture
of local environment at the LGM/LGT
in the context of the Middle Danube region
Zdeňka Nerudová,1 Nela Doláková2 and Jan Novák3
Abstract
Records of occupation by humans in the period following shortly after the Last Glacial Maximum (LGM; 21 ± 2 kyr cal. BP) are still very rare in Central
Europe, since it is inferred that the extreme climatic conditions caused the decolonisation of previously settled areas. Our study focuses on the
reconstruction of environmental conditions in the surroundings of the open-air Palaeolithic site, Brno-Štýřice III, which falls within this period. The
research concentrated on the study of malacological, pollen and anthracological samples to reconstruct the climate shortly after the LGM. 14C dating
places the chronostratigraphic position of the site more precisely at the end of the LGM, more specifically into Last Glacial Termination (LGT); analysis of
chipped stone industry identifies the occupation with the Epigravettian settlement. The site represents a significant example of the recurrent habitation of
a microclimatically favourable microregion near a watercourse in order to utilise available sources of livelihood. The results of the pollen, anthracological
and malacological analyses documented a more or less treeless character of surrounding landscape. The vegetation was mostly formed by a mixture of
shrub tundra and grassy loess steppe vegetation. Open woodland with birch, willow and bird cherry occurred in relatively moist river banks and the lower
slopes of hills with more favourable microclimatic conditions. Malacological collection highlights the presence of cool temperate species (Pupilla loessica,
Vallonia excentrica and Helicopsis striata). In the surroundings of the studied site, the pollen analysis provided a reconstruction of parkland forest-steppe
vegetation (with lack of temperate deciduous trees) typical for a cold and dry climate. Development of both dry and moist stands near the watercourse
was recorded. Anthracological analysis is in support of similar outcomes, reconstructing the presence of open woodland with dominating birch and willow
in the nearby surroundings.
Keywords
Anthracology, LGM/LGT, malacology, Middle Danube region, Palaeolithic settlement, palynology
Received 12 November 2015; revised manuscript accepted 22 February 2016
1Moravian Museum, Historical Museum, Centre for Cultural
Anthropology, Czech Republic
2Institute of Geological Sciences, Faculty of Science, Masaryk University,
Czech Republic
3Laboratory of Archaeobotany and Palaeoecology, Faculty of Science,
University of South Bohemia, Czech Republic
Corresponding author:
Zdeňka Nerudová, Moravian Museum, Historical Museum, Centre for
Cultural Anthropology, Zelný trh 6, Brno, CZ-65937, Czech Republic.
Email: znerudova@mzm.cz
640051HOL0010.1177/0959683616640051The HoloceneNerudová et al.
research-article2016
Research paper
1346	 The Holocene 26(9)
Willis et al., 2000). Nevertheless, palaeobotanical data from LGM
sites in eastern-central Europe generally confirm the existence of
not only parkland landscapes with coniferous trees (such as Pinus
sylvestris, Pinus cembra, Larix, Picea and Juniperus) but also
more demanding tree taxa (Abies, Corylus, Quercus, Fagus,
Fraxinus, Ulmus, Taxus or Carpinus; see Jankovská and Pokorný,
2008; Willis and Van Andel, 2004). Taking into account new palaeobotanical
and molluscs data, the local conditions in the whole
area of the Czech Republic were more favourable (for more discussion,
see Horsák et al., 2010; Jankovská and Pokorný, 2008;
Juřičková et al., 2014; Ložek, 2001, 2009). The aim of this study
is to present the new information about the local climatic conditions
during the LGM/LGT period based on the results of excavation
of a Palaeolithic site.
Archaeological background
From the archaeological point of view, the LGM period in Central
Europe is characterised by the end of Gravettian in the so-called
Kostenki-Willendorf phase (around 24–25 kyr cal. BP) and the
onset of Magdalenian colonisation, which proceeded from Western
Europe through the north German lowlands as far as Moravia,
where it arrived in around 17–18 kyr cal. BP (Neruda, 2010; Neruda
et al., 2009). Despite the generally adopted awareness of the
decolonisation of the climatically exposed regions of Central
Europe in the LGM and the following period that preceded the
Magdalenian colonisation, in recent years, we applied ourselves
to the study of an archaeological site, the occupation of which
falls within the period, when the LGM petered out, and the LGT
that followed, on the grounds of 14C dating (Nerudová and Neruda,
2015). The evidence of the local more favourable conditions
can be documented not only by palaeorecords in vegetation but
also by archaeological evidence. On the basis of the cultural classification
or geomorphological position we can associate with the
short period between the end of the Gravettian and before the
onset of the Magdalenian in Moravia almost 49 sites. Contrary to
the previous Gravettian, the re-analyses of the settlement dynamics
revealed significant changes. On the basis of reconstruction of
settlement strategy of the Epigravettian sites we can conclude that
the majority of localities are situated within the interval of 200–
250 m a.s.l.; the sites are mostly oriented towards the east through
to north-east and are usually much closer to (mostly) small river
in protected areas (Nerudová and Neruda, 2015).
Regional setting, site
characteristics and stratigraphy
The excavation site Brno-Štýřice III (global positioning system
(GPS): World Geodetic System 1984 (WGS-84): 49°11′2.5505″N,
16°35′41.6602″E; S-JTSK: 1161873.78, 599243.33 – the centre
of the locality) is located in the south-western part of Brno,
approximately 300 m to the south of the current bank of the
Svratka River (Figure 1). Here, at an elevation of 210 m a.s.l.
(10 m above the river level) is a step in the lower terrain which on
the west side rises up into a low but steep cliff of Lower Devonian
conglomerates known as Červený kopec (Red Hill) with a maximum
height of 311.42 m a.s.l. The Quaternary cover of the region
is formed by an accumulation of aeolian (loess) and colluvial
sediments deposited on a terrace consisting of gravels with clay
matrix and sandy gravels of Quaternary age, which were detected
at a depth of 6–8 m. A coring revealed that the sequence of Pleistocene
sediments at the locality is up to 10 m thick in places and
is not divided by any distinct fossil soils. Towards the superposed
layers, the sequence of loess and loess sediments is covered by an
orange silty sediment C (weakly developed soil followed by
Holocene stratigraphy brown earth; see Nerudová et al., 2012:
Figure 2). Within the whole area under investigation, Holocene
soil inclusive of the A- and B-horizons is only preserved in higher
parts (i.e. at the W and SW edge of the excavated surface). In the
central part of the slope, only a relic of the B-horizon is preserved,
and in the NE and E parts of the investigated area (i.e. in the lowest
parts), the B-horizon is not preserved at all. This being the
reason why the A-horizon settled immediately on the Pleistocene
sediments.
The site of Brno-Štýřice III was first investigated by K Valoch;
in 1972, he conducted small-scale rescue excavations there; nevertheless,
the excavations yielded a representative collection of
chipped stone industry as well as the remains of animal bones and
blobs of ochre (Valoch, 1975). The assemblage was dated later on
(Valoch, 1980, 1996; Verpoorte, 2004: 262); for more discussion,
see Nerudová and Neruda (2014). The next large-scale rescue excavations
were carried out in 2009 and 2011–2014. They revealed the
extent of settlement, a new site (in 2009 – Štýřice IIIa), and yielded
a large amount of lithics and osteological material.
Archaeological finds of the Palaeolithic age were found in the
uppermost part of the Late Weichselian loess cover, which formed
a 25-cm-thick (approximately) layer of orange-brown loess-like
sediment (weakly developed soil). This layer was almost continuously
present over the entire investigated area as it followed the
inclination of the terrain, which was quite steep in certain places.
There was a relatively sharp border between the horizon with
archaeological finds and the underlying sediments. Although the
lithic artefacts and bones were deposited in loess-like sediment,
the base of the superposed chernozem horizon A sporadically
intruded (e.g. the upper part of a mammoth jawbone; Nerudová
et al., 2012). The first radiocarbon date was obtained from a
charred bone of reindeer (Valoch, 1980, 1996), the next dating
(performed by A Verpoorte in 2001 from the same sample; see
Nerudová and Neruda, 2014: Table 4), yielded a date very similar
to the previous one (Verpoorte, 2004). The evidence of human
activities falling the site within the Epigravettian period (Nerudová,
2015; Valoch, 1975). Sampling during the last seasons of
excavations provides us to perform the palaeobotanical analysis.
Materials and methods
The collection of samples
The collection of samples was carried out within the archaeological
research of the site in 2009 and 2011–2014 (Figure 2; e.g.
Nerudová and Neruda, 2014). We took samples of varying volumes:
individual bigger charcoals (0.5–2 cm) macroscopically
distinguishable in the course of preparation of the archaeological
layer (all bigger samples taken during excavation were determined
as fragments of burnt bones; Roblíčková et al., 2015) and
target sediment samples of 5–10 L from the places with dispersed
pieces of charcoal or bones. For the pollen analysis, we took samples
of 0.5–1 L volume.
The extractions of the charcoal, molluscs and bones from the
sediment samples were subjected to the standard flotation procedure
using staggered sieves with a mesh size of 0.25 mm.
Radiocarbon dating
New radiocarbon dating was performed using the accelerator
mass spectrometry (AMS) method in the laboratory in Oxford,
where samples underwent standard laboratory procedure (Bronk
Ramsey et al., 2004b). Animal bones (mammoth molars and
mandible) coming from a distinct archaeological context served
for the dating because of the volume required, since no other
suitable material was available (the abundance and size of charcoals
was very low, at the limit of the method; see Bronk Ramsey
et al., 2004a). Before dating, the bones were not cleaned using
chemical methods; they were only determined. The standard process
in Oxlab is measurement of bone collagen by ultrafiltration
Nerudová et al.	 1347
method (Bronk Ramsey et al., 2004b). Isotopic fractionation has
been corrected for using the measured 13C values measured on
the AMS, the quoted 13C values are measured independently on
a stable isotope mass spectrometer (see Bronk Ramsey et al.,
2004a, 2004b).
Laboratory preparations and analysis of fossil
samples
Palynological samples were picked from squares 83 and 95 (two
samples from each of them). An accumulation of charred bones
and overburnt sediment was detected in square 95, consequently
a sediment was taken for palynological analysis; unfortunately,
the pollen record from square 95 was very poor (see Table 2).
Palynological samples were treated with HCl (20%), HF, KOH
and HCl (10%) and heavy liquid ZnCl2 (density = 2 g/cm3) for
standard maceration. The omission of acetolyse enabled clearer
identification of contemporary pollen contamination. The final
residue was diluted with glycerol. A Nikon Alphaphot 2 light
microscope with 400× and 1000× magnifications was used for
palynological observations. Taxa identification was mainly
done according to Reille (1995) and Beug (2004). The pollen
diagram was created using the POLPAL programme (Walanus
and Nalepka, 1999).
Figure 1.  (a) Moravia region with LGM/LGT sites mentioned in the text and (b) microregion of the Brno-Štýřice with Palaeolithic finds.
1: Štýřice III; 2: Štýřice IIIa; 3: Kamenná St; 4: Hospital; 5: Polní St; 6:Vídeňská St no. 15; squares: isolated or sporadic finds; circles: extent sites.
Source: Digitalisation – Z. Nerudová.
1348	 The Holocene 26(9)
The charcoal analysis was performed only on fragments from
the largest fraction (>1 mm). The charcoals were identified using
an episcopic interference microscope (Nikon Eclipse 80i) with
200–500 magnification and the reference collection. The additional
standard identification keys were also used (Heiss, 2000;
Schweingruber, 1990). The species abundance was expressed in
the number of charcoal fragments (e.g. proposed by Delhon,
2006) and charcoal anthracomass (e.g. Carcaillet and Thinon,
1996). The individual taxa were weighted with an accuracy of
0.001 g. The sediment anthracomass (milligram of charcoal per
kilogram of sediment; Talon et al., 1998) was derived from the
charcoals larger than 1 mm.
Mollusc shells for malacological studies were derived from
individual sediment samples taken with the bone samples for
laboratory processing.
Results
Radiocarbon dating
Most recent radiocarbon dating was done on samples taken during
the excavations in 2009 and 2011. Only a single date from a mammoth
molar was obtained because of the low collagen content of
the other samples. Dating of other samples from the 2012 excavation
was successful. From a total of four samples, only two dates
were obtained, and these dates were similar to the previous ones
(Table 1). Sample of burnt bone (OxA-28114) gave a good yield of
carbon on combustion, and a CN ratio of 7, which indicates that
there is some pyrolysed collagen remaining. In addition, the stable
isotope ratio (d13C) was −22.2, which is acceptable for a large
mammal. For this reason, the sample was OxA’ed. Dating of the
next two samples of charred bones taken from the excavation in
2014 unfortunately failed because of very low yield.
Pollen analysis
A very interesting pollen spectrum was found in samples from
square 83. Predictably about one-fifth of the total number of the
determined pollen grains and spores consisted of palynomorphs
redeposited from the older Tertiary sediments. Woody plants
were relatively amply represented by pollen grains of pine with a
slight prevalence of common Pinus sylvestris over Pinus cembra.
Other species that were determined include Betula – finding
of three grains in a clump testifies to a short transport, less commonly
Alnus and sporadically Corylus (Figure 3). From the
pollen grain ratios of woody species (arboreal pollen (AP)) 52%
Figure 2.  Detail plan of the Brno-Štýřice III site (excavations
2009–2014) and positions of taken samples for analyses.
Black dot: pollen sample; asterisk: molluscs; square: charcoals.
Source: Digitalisation – Z. Nerudová.
Table 2.  List and abundances of pollen grains.
Square 83 Square 95
Arboreal pollen (AP)
 Alnus 2  
 Betula 6  
 Corylus 1  
 Ephedra 1  
  Pinus sylvestris 35  
  Pinus cembra 24  
  Rosaceae type Rubus 1 1
Nonarboreal pollen (NAP)
 Artemisia 1  
  Asteraceae, Cichorioideae 4  
  Asteraceae,Asteroideae 5 2
  Brassicaceae type Cardamine 1 1
 Chenopodiaceae 6 1
 Chrysosplenium 6  
 Cyperaceae 2  
 Daucaceae 1  
 Ericaceae 1  
 Helianthemum 1  
 Persicaria 1  
 Plantago 1  
 Poaceae 10 1
 Glyceria type 2  
 Ranunculaceae 3  
  Caltha 2  
  Delphinium 4  
 Thalictrum/Illecebrum 5  
 Veronica 3  
Sporophyta
 Botrychium 1  
 Lycopodium 1  
  Polypodiaceae smooth 1  
 Sphagnum 1  
 Bryophyta 2  
Algae
  Botryococcus braunii 16 1
 Pediastrum 1  
 Circulisporis 6  
Fungi xx x
 Glomus 3  
Tertiary redepositions undetermined 21  
Table 1.  14C data from Brno-Štýřice III.
OxA-26961: 15,625 ± 75 BP (excavation 2009; molar, Mammuthus
primigenius)
OxA-28298: 15,215 ± 70 BP (excavation 2012; molar, Mammuthus
primigenius)
OxA-28114: 14,870 ± 90 BP (excavation 2012; charred bone, Mammuthus
primigenius)
Nerudová et al.	 1349
and herbs (nonarboreal pollen (NAP)) 48%, the character of vegetation
appears to be a moderately forested landscape to parkland
(Table 2). However, the proportion of woody plants (in which
pollen of Pinus markedly prevails) can be greatly overvalued
because of prodigious pollen production and the great flying
range of pine pollen grains. The herbal constituent of the spectrum
included the representatives of both drier and waterlogged
habitats; for example, grasses Poaceae, floscular Asteraceae,
Artemisia, or Chenopodiaceae occurred in drier areas, ascertained
were Delphinium or Veronica. Bushes such as Ephedra,
Rubus type or Helianthemum and Ericaceae were also represented.
Elements characteristic for damp, waterlogged habitats
or watersides such as Cyperaceae, Glyceria, Caltha, Chrysosplenium
or scarce spores of Sphagnum were also found. Colonies
of aquatic Botryococcus braunii were relatively abundant;
those of the Pediastrum were sporadic. A paucity of thermophilic
woody species together with taxa typical for cold periods of the
Quaternary such as Helianthemum, Thalictrum and Ephedra
determines the character of the climate.
Anthracological analysis
A total of 89 charcoal fragments and 6 charcoal species from 13
samples were identified. Anthracological samples were distinguished
by the low presence of small pieces of charcoals (the
most common charcoal size was 2–3 mm). The anthracomass of
analysed samples was very low. A significant feature of the entire
assemblage of samples was a considerable prevalence of carbonised
bone fragments over wood charcoals.
In the charcoal samples obtained from the Epigravettian layer,
the dominant tree was Betula (34.8%), followed by abundant
charcoal of Salix (27.0%) and Padus (21.3%). The presence of
Picea/Larix (11.2%) was relatively common. Only scarce occurrences
of Hippophae (4.5%) and Ericaceae (1.1 %) were recorded
(Figure 4).
Malacological analysis
The first representatives of malacofauna that were found are
Pupilla loessica (9 pcs), Vallonia excentrica (1 pc) and Helicopsis
striata (1 pc) coming from the hearth area (square 9/Q, excavation
2009; determination by L. Juřičková; Nerudová et al., 2012).
Other individuals of the Pupilla loessica species (determination
by M. Horsák) were found during the next season of excavation
(i.e. 2012): square 35 (6 pcs), square 48 (2 pcs), square 49 (1 pc),
square 26 (1 pc) and square 22 (1 pc).
Figure 3.  Photographs of the main pollen types (square 83), magnification 1000×.
1: Pinus cembra type; 2: Betula – three grains; 3: Helianthemum; 4: Rosaceae type Rubus; 5: Ephedra; 6: Glyceria type; 7: Thalictrum; 8: Caltha.
Source: Photograph – N. Doláková.
Figure 4.  Number of charcoals in the study samples.
1350	 The Holocene 26(9)
Discussion
Proxy data serving for the reconstruction of palaeoclimate could
not be obtained through systematic sampling, because neither the
state of preservation and the character of the Pleistocene sediments
nor the character of rescue archaeological excavations
allowed it. Despite several pollen samples taken, only one was
positive for determination, and in this single case, it was taken
close below the archaeological layer level (at its base). Relatively
numerous shells of bivalves were found solely in relation to laboratory
processing of large fragments, especially mammoth bones;
at the same time, it has to be stressed that microfauna was not
ascertained at all. In view of the soil chemistry influenced by
post-deposition processes, the shells were always preserved only
in the sediment and close to scarce bigger bone fragments or
hearths; contrary to the surrounding area, such environment is
always more calciferous, which facilitated their preservation
(Nerudová et al., 2012; environmental requirement in general, for
example, in Juřičková et al. (2014)). Determination of the osteological
material was greatly impeded by its fragmentary character
and severe surface deterioration, even in bones of large mammals
with thick compact bones (Roblíčková et al., 2015). However,
although we took all of the macroscopically apparent charcoals,
and also the entire sediment from the places of hearths, the numerous
tiny burnt fragments turned out to be animal bones, not woody
species. Nevertheless, both archaeological and palaeobotanical
analyses brought interesting, mutually complementary and corresponding
results that are in harmony with the radiocarbon dating
of the site.
Pollen analysis indicates scarcity of temperate woody species
at the site and the presence of taxa typical for cold periods of the
Quaternary determining the character of the climate as cold and
dry. Parkland vegetation with the development of both drier and
waterlogged habitats near water streams was reconstructed.
According to the determination of malacozoological analysis,
the ascertained individuals belong to the typical representatives
of loess steppe, since in the late glacial Pupilla loessica
fades away (Horáčková et al., 2015; Horsák et al., 2010; Ložek,
1990, 2001, 2006).
Anthracological analysis recorded a dominance of Betula,
common presence of Salix, Padus, Picea/Larix and only scarce
occurrence of Hippophae and Ericaceae in study samples. These
results indicate a cold, dry climate which is consistent with results
and reconstructions. The matrix of landscape was probably more
or less treeless. Open woodland with Betula, Salix, Padus and
Picea/Larix was restricted to relatively moist banks of the river
and protected valley. We can find the recent vegetation analogy
in the river banks in southern Siberia (Magyari et al., 2014a) or
northern Mongolia. The mixture of Salix, Padus, Betula (e.g.
Betula fusca), Picea obovata and Hippophae rhamnoides formed
typical vegetation composition in the vicinity of rivers or streams.
The cold and dry climate at the site is also indicated by the
representatives of mammoth steppe fauna (of which most plentiful
was exactly the Mammuthus primigenius, with Equus
germanicus, Rangifer tarandus or Coleodonta antiquitatis as
accessory species) and the results of analyses of carbon and nitrogen
isotopes. Numerous fragments of animal bones could have
resulted from deliberate human activity, substituting bones for
shortage of wood in a period of shortage during the glacial maximum.
Similar practices were ascertained, for instance, in
Předmostí I-06 (Beresford-Jones et al., 2010), Dolní Věstonice II
(Beresford-Jones et al., 2010; Svoboda, 1991a) or GrubKranawetberg
(Bosch et al., 2012), and researches show the use
of bones as fuel was a ‘common behavioural pattern during the
Middle and Upper Palaeolithic in Northern Europe’. The use of
bones as fuel was described in detail in the contribution by Bosch
et al. (2012) since a more detailed analysis of this phenomenon is
not a subject of this study.
This is where the results of radiocarbon dating and other scientific
analyses diverge to some degree. After calibration (CalPal:
Weninger and Jöris, 2008, and IntCal2014: Reimer et al., 2013),
all of the data acquired from the site of Brno-Štýřice III so far
form two peaks with an interval of roughly 1000 years GrN 9350:
17,620 ± 120 cal. BP and OxA 26961: 18,880 ± 90 cal. BP, that is,
just after the stated interval of LGM (21 ± 2 kyr cal. BP). All new
14C dates were obtained from the mammoth bones and mandibles,
representing the only material found at the site from which it was
possible to at least partly obtain any dates; these bones were
found in close proximity to stone artefacts and within the same
archaeological layer. Tools were also found in the sediment within
the mandibles. We consider this close contextual association to be
strong presumptive evidence for the contemporaneity of the lithic
industry and mammoth bones, and considering the homogeneity
of the dates, it is very possible that the site’s occupants gathered
the remains of mammoth carcasses.
However, the results of the pollen, anthracological and osteological
analyses documented a similar character of surrounding
habitats. The landscape was more or less treeless. The vegetation
was mostly formed by shrub tundra vegetation with grassy loess
steppe. Open woodland with birch and willow occurred in relatively
moist river banks (of the nearest Svratka River) and foot of
a hill (the Červený kopec Hill) with more favourable microclimatic
conditions (Ložek, 2001, 2009).
V. Ložek (2010) states,
loess provides most evidence on the glacial environments in
dry warm areas of Central Europe, which – in contrast to the
conditions in north-western Europe – … never had the character
of northern tundra, but of cold continental steppes and
barren lands of InnerAsia instead.Aclear testimony brings the
loess fossil fauna, primarily the almost ubiquitous snails, the
communities of which were preserved in the loess in secondarily
undisturbed original appearance, enabling a critical comparison
with similar recent snail assemblages of Inner Asia.
Nevertheless, on the grounds of our environmental analyses,
manifestations of a slight warming in this period were not corroborated.
Considering that the improvement of climatic conditions
in the period shortly after the LGM (LGT) was only
moderate, the changes in vegetation and fauna were probably
relatively small; the microclimatic factor that enables surviving of
typical zoocenoses (e.g. snails) in microclimatically suitable conditions
in the face of the generally prevailing climate has to be
considered as well (e.g. Horsák et al., 2015; Juřičková et al.,
2014; Ložek, 1990, 2001, 2006).
Regretfully at present, we do not have larger quantity of data
regarding the LGT environment in archaeological context available
for the studied region. Therefore, we have to rely on information
characterising the previous last cold oscillation instead.
If we take into account published isotopic data for mammoth
in Europe and results of analyses of carbon and nitrogen isotopes
in Brno-Štýřice III (Roblíčková et al., 2015), the results fit well
together and generally confirm a cool steppe environment with
low precipitation (Kovács et al., 2012; Pryor et al., 2013). The
isotopic values from Brno-Štýřice III correspond to the Late Gravettian
environment, especially in Moravia region (Roblíčková
et al., 2015).
For the Middle Danube region in the LGM period, the landscape
is characterised as a combination of cold loess steppe and
mosaic parkland (Feurdean et al., 2014; Heiri et al., 2014). In central
and eastern part of Europe, the major vegetation type of megabioms
of the type cold deciduous trees is represented by Alnus,
Betula, Salix and Populus. Picea, Pinus, Abies, Larix and Juniperus
represent coniferous trees. Grass and shrubs are represented by
Nerudová et al.	 1351
Ericaceae, Calluna, Hippophae, Poaceae, Cyperaceae and other
NAPs. Artemisia and Chenopodiaceae/Amaranthaceae represent
xerophytic herbs (Feurdean et al., 2014; Heiri et al., 2014).
We have to consider the station at Stránská skála IV the nearest
analogous and at the same time archaeological site. The Epigravettian
site, dated at 18,220 ± 120 and 17,740 uncal. yr BP, yielded
evidence of hunting of Equus, which prevailed, Rangifer tarandus,
Bos, Mammuthus and Coleodonta antiquitatis. The mollusc successions
acquired is indicative of a major biological rearrangement
of the sediment since together with Pupilla loessica, also a striatic
fauna was found, if it were a homogeneous successions (Svoboda,
1991b). Herb assemblage was also acquired from this site, and
stratigraphy made its assessment complicated (the Epigravettian
industry was situated in the uppermost part of the Weichselian
loess also in this area; thus, it became secondarily turned into soil
by the superposed Holocene horizon). Together with less climatically
demanding woody species (Betula, Salix, Pinus, Corylus and
Alnus), the sample also contained herbs (Cyperaceae, Poaceae,
Artemisia, Asteraceae-Tubiflorae, Ericaceae, Lythrum, Chenopodiaceae
and Silenaceae), ferns (Polypodiaceae) and moss (Sphagnum;
Svoboda, 1991b).
In the South Moravian region, the profiles in Dolní Věstonice
II or in Bulhary can be considered the closest regional and temporal
analogous situation. Pollen analyses indicate that towards
the end of the Weichselian interpleniglacial, coniferous trees
(pine and spruce) were prevalent in Dolní Věstonice II, although
more demanding woody species such as oak and beech also
appear sporadically. Pollen grains of these more demanding species
were also ascertained in the filling of the calva from triple
burial at this site. According to H. Svobodová (2002), the herb
spectrum points to forest–steppe character of the environment
(Svobodová, 1991). Water and peat environment in the surroundings
is indicated by Myriophyllum, Sparganium, Potamogeton,
Trapa natans and algae (Svobodová, 1991). Anthracological
analysis supports the presence of Larix, Picea/Larix, Pinus sylvestris,
Pinus cembra, Betula, Hippophae and Juniperus at this
site (Svoboda et al., 2015).
Anthracological analyses from the nearby Dolní Věstonice I
confirmed the occurrence of pine, spruce and larch; further analyses
testify to fir, perhaps also poplar (see Beresford-Jones et al.,
2010; Svobodová, 2002). In the pollen spectrum, obtained from
the core situated near the Bulhary Village and reaching down to
about 10 m of aeolic loess layers and some 40–50 cm in thickness
of compressed organogenic sediments (the moss peat and algal
gyttja; 25,675 + 2.750–2.045 BP; Hv 10,855), coniferous species
prevailed, birch, juniper, but also scattered temperate deciduous
trees (Ulmus, Acer, Quercus and Tilia) were found, and these
could grow at favourable sites in relation to relief. Unfortunately,
the result of pollen analysis from this locality reconstructed vegetation
closely before LGM. In the assemblages, over 200 types
of palynomorphs (pollen, spores, etc.) have been found, which
indicates a very rich flora and vegetation growing in at least six
different biotopes of the Pálava Hills region (Rybníčková and
Rybníček, 2014). The spectrum indicates grass and herb plant
steppe community. The vegetation also comprised aquatic and
swamp plants and ferns (Rybníčková and Rybníček, 1991, 2014;
Svoboda et al. 2002).
Not only anthracological or pollen results but also various
malacological analyses corroborate that the sporadic but several
times observed findings of temperate deciduous trees could indicate
the existence of refugia, which were preserved in more
favourable enclaves – on southerly oriented slopes of the Pálava
Hills – after the climatic deterioration (Juřičková et al., 2014;
Ložek, 2006, 2009). According to Musil (1999, 2003), we can
presume that such refuge areas facilitating growth of woody species
requiring higher temperatures could have persisted since the
previous interglacial; for the area of Moravia, Ložek (2009) or
Horáčková et al. (2015) and Juřičková et al. (2014) argue for their
existence on the grounds of malacozoological studies.
We know of pollen spectra capturing the LGM period from
several sites in the Moravian Karst, for example, the Kůlna (layer
6; see Svobodová, 1988) or Barová (Seitl et al., 1986) and the
Balcarka Caves (Doláková, 2010). At Balcarka Cave, prevailing
herbs were found together with pollen of Betula in the sediment
dated at 28,360 uncal. BP, captured inside the cave (Doláková,
2010). Within layers 11 and 12 in the Barová Cave (Seitl et al.,
1986), the share of woody species increased and decreased several
times (amounting up to 56% in the central part) – with prevalent
Pinus, present Betula, Alnus, Picea and Corylus. In the herb spectrum,
Poaceae and Asteraceae prevailed; Helianthemum, Selaginella,
and Ephedra were present. Broadleaved trees requiring
higher temperatures were not captured at the site. Nevertheless,
alder, spruce and hazel do not represent typical tundra vegetation,
either. Watercourses and the rugged relief (compare Lisá et al.,
2013) probably made the climate more moderate in these areas.
The presence of macrofossil and charcoal records from the
period shortly after LGM is still very rare in Central Europe. In
addition to the presented site of Brno-Štýřice III, another significant
site in the broader regional context is Mohelno-Plevovce
(Škrdla et al., 2015); upon the anthracological analysis, it reconstructs
the vegetation shortly (19,690 ± 120 cal. BP) after the
LGM. The results of anthracological analysis indicate cold
and dry shrub tundra vegetation, patches of open woodland with
Salix, Betula and Juniperus and grassy loess steppe (Škrdla
et al., 2015).
Similar climatic and vegetation conditions, closely compared
with the Central Europe LGM climate, were described in detail
from Altai region in southern Siberia. This region is recently
regarded as the best known modern analogue of the last Pleniglacial
of central Europe (Horsák et al., 2010). The results of studied
spectrum of snail taxa indicates, that the full-glacial landscapes of
Central European lowlands were not completely dominated only
by open and dry loess steppe, but they contained a significant
component of shrubby vegetation, patches of wet habitats and
probably also areas of woodland at sites with a more favourable
climate (Horsák et al., 2010).
Conclusion
At Brno-Štýřice III, evidence of human occupation in the end
of the last (Weichselian) glacial was situated in the uppermost
part of the Last Weichselian loess affected by intense Holocene
processes of turning into soil. As a typical aeolian calcareous
sediment, loess facilitates preservation of a certain type of material,
especially terrestrial snails.
Despite unfavourable deposition conditions, we managed to
ascertain new information augmenting the picture of local environment
of the LGM/LGT in the context of the Middle Danube
region. Although this information confirms the contemporaneous
incidence of plant assemblages favouring lower temperatures, at
the same time, it is evident that humans chose suitable temperate
refuges close to water resources for their settlement, where they
were adequately suggest more favourable microclimatic conditions
protected from extremes of the climate.
The ascertained species composition of anthracological samples
differs from the collections from other Central European
LGM sites (e.g. Dolní Věstonice IIa; Svoboda et al., 2015); this
may be influenced by the position and character of settlement.
Abundant presence of charcoals of Salix and Padus corresponds
to microclimatically more favourable situation of the site near a
watercourse. Scanty representation of woody species charcoals
against very abundant presence of charcoals from bones is indirect
evidence of scarcely represented woody species in the vicinity
of the studied site, whereas it has to be mentioned that low
1352	 The Holocene 26(9)
abundance of charcoal was probably recorded also at other
archaeological sites dated back to the LGM period (Předmostí).
The charcoal analysis reconstructed the presence of sparse
birch groves in the vicinity. These groves were probably tied to
microclimatically favourable conditions (e.g. damper northern
slopes) within the prevailing cold steppe or steppe–tundra.
Recently, it is possible to come across analogous types of vegetation
in Siberia (Horsák et al., 2010, 2015; Magyari et al., 2014a) or
Mongolia. These habitats are characterised by both low temperatures
during winter season and cold dry winds and very low rainfall,
which impose large limitations on the presence of dense
coniferous forests (dark tajga forests). Numerous fragments of
animal bones could have resulted from deliberate human activity.
We would like to point out the importance of the results, especially
in the context of the still poor known archaeological background,
which is based mainly on the nonstratified sites. At
Brno-Štýřice III, occupation by humans and species composition
of vegetation was apparently influenced by the mentioned vicinity
of a large waterway. Keeping to the outcomes of radiocarbon
dating together with other analyses, we draw a temporal link of
the settlement in Brno-Štýřice III more probably with the LGT
period. The site is an exemplification of a microclimatically
favourable refuge area serving for humans towards the end of
Upper Palaeolithic, the period for which absence of archaeological
evidence in a considerable part of Europe corroborates the
decolonisation of vast areas prior to the onset of the Magdalenian
settlement. We hope that the results will be helpful in the discussion
concerning the climatic conditions and LGM/LGT central
European occupation.
Acknowledgements
We are very grateful to Michal Horsák and Lucie Juřičková for
determination of mollusc fauna and Martina Roblíčková for many
valuable comments on the faunal composition. We would like to
mention the anonymous reviewers for their valuable comments
on this manuscript.
Funding
The research reported here was financially supported by the Ministry
of Culture of the Czech Republic by institutional financing
of long-term conceptual development of the research institution
(the Moravian Museum, MK000094862; Z. Nerudová), institutional
support of Institute of Geological Sciences, Masaryk University
– 312222 (N. Doláková) and GAČR 13-11193S (J.
Novák).
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