Geo log i cal Quar terly, 2016, 60 (2): 461–472 DOI: http://dx.doi.org/10.7306/gq.1289
Homotryblium-dom i nated Eocene dinoflagellate cyst as sem blages
from Mid dle Mio cene (Badenian) glauconitic sands at Lipowiec (Roztocze, SE Po land)
Przemys³aw GEDL1, *
1 Pol ish Acad emy of Sci ences, In sti tute of Geo log i cal Sci ences, Senacka 1, 31-002 Kraków, Po land
Gedl, P., 2016. Homotryblium-dom i nated Eocene dinoflagellate cyst as sem blages from Mid dle Mio cene (Badenian) glauconitic sands at Lipowiec (Roztocze, SE Po land). Geo log i cal Quar terly, 60 (2): 461–472, doi: 10.7306/gq.1289 A sec tion over 20 m thick of the basal Mid dle Mio cene suc ces sion ex posed at Lipowiec (Roztocze, SE Po land) was stud ied for palynology. Thir teen sam ples were col lected from quartz and glauconitic-quartz sands and thin clay lay ers. Sand sam - ples were bar ren but clay sam ples yielded dinoflagellate cysts. Their as sem blages con sist of re worked Paleogene spec i - mens dom i nated by the Homotryblium floripes com plex (H. floripes and mor pho log i cally sim i lar H. plectilum and H. vallum).
The pres ence of re worked Paleogene spec i mens in di cates in tense ero sion of ma rine Paleogene strata dur ing ini tial stages of the Mid dle Mio cene trans gres sion at Roztocze. Anal y sis of strati graphi cal ranges of re worked dinoflagellate cysts and com par i son with their oc cur rences in known epicontinental Paleogene sites of SE Po land sug gest an Up per Eocene age of the washed-out strata. The tax o nomic com po si tion of the as sem blages de scribed sug gests that the Up per Eocene de pos its ac cu mu lated in a near-shore, la goonal embayment char ac ter ized pre sum ably by oligotrophic wa ters of slightly in creased sa - lin ity. A high pro por tion of the Homotryblium floripes com plex is also char ac ter is tic of re worked as sem blages found in youn - ger Mio cene strata of neigh bour ing ex po sures in Roztocze, which in di cates wide spread Eocene la goonal en vi ron ments in this part of Roztocze. Co eval Eocene strata from the east ern part of Roztocze and from the Carpathian Foredeep show dif fer - ent tax o nomic com po si tions sug gest ing var ied sed i men tary set tings dur ing Eocene trans gres sion in this re gion. These dif - fer ences re flect vari able amounts of fresh wa ter in flu ence re sult ing in a range of en vi ron ments that ranged from oligotrophic to brack ish.
Key words: Eocene, Mio cene, re work ing, palaeoenvironment, Roztocze, SE Po land.
INTRODUCTION
Eocene and Oligocene ma rine strata were for merly widely dis trib uted in SE Po land cov er ing a part of the Carpathian fore - land that now forms the Roztocze hills and the east ern part of the Carpathian Foredeep. Most of them were re moved by post-Rupelian ero sion that left only a few sites with pre served de pos its of these ages (e.g., Buraczyñski and Krzowski, 1994;
GaŸdzicka, 1994; Gedl, 2000, 2012, 2014, 2015; Myœliwiec and Œmist, 2006). This makes re con struc tion of Eocene and Oligo - cene palaeo ge ogra phy in SE Po land, and of their palaeo - environmental char ac ter is tics, dif fi cult. In such cases, re worked Paleogene microfossils, dinoflagellate cysts in par tic u lar, which oc cur in Mio cene strata of the Carpathian Foredeep, are help - ful. Re worked dinoflagellate cysts have been de scribed from sev eral Mio cene sec tions, in di cat ing a much wider ex tent of Eocene and Oligocene epicontinental seas in SE Po land than could be in ferred from the ba sis of pre served Paleogene sites (Gedl, 2012).
Mid dle Mio cene suc ces sion of Roztocze ap pears to be par - tic u larly rich in re worked Paleogene, par tic u larly Eocene, dino - flagellate cyst spe cies. Fre quent re worked forms, com monly out num ber ing Mio cene spec i mens, have been de scribed from Up per Badenian de pos its that rest upon the evaporitic ho ri zon at Józefów (Gedl, 2012, 2016).
Fur ther dis cov er ies of re worked Paleogene dinoflagellate cysts in the Lower Badenian strata of Roztocze, pre sented in this pa per, in di cate ex pan sion of the time in ter val dur ing which the Paleogene cover un der went ero sion. I pro vide strati graphi - cal anal y sis of the as sem blages to al low their dat ing, and qual i - ta tive anal y sis to make palaeoenvironmental re con struc tions of now non-existing Eocene strata.
GEOLOGICAL BACKGROUND
The Carpathian Foredeep Ba sin de vel oped as a fore land ba sin at the front of the north wards-mi grat ing Carpathian orogen. It stretches along the front of Carpathians from Aus tria to Ro ma nia, a dis tance of over 1300 km. In the Pol ish part (Fig.
1A), the Carpathian Foredeep is di vided into in ner and outer parts. The in ner part is now a days cov ered by overthrusted Carpathian nappes, or oc curs as nar row belts of folded strata at the front of the Carpathians. It in cludes the Lower (mainly con ti - nen tal) and Mid dle Mio cene (Badenian and Sarmatian de pos -
* E-mail: ndgedl@cyf-kr.edu.pl
Received: February 11, 2016; accepted: March 30, 2016; first published online: April 11, 2016
its), which are ma rine. The outer part of the Carpathian Foredeep is filled with Badenian and Sarmatian ma rine strata that reach up to 2000 m in thick ness, and 5000 m in the Ukrai - nian part (for de tails see e.g., Ney et al., 1974; Kotlarczyk, 1985; Oszczypko et al., 2006).
The Mio cene se quence of the outer part of the Carpathian Foredeep is as so ci ated with a ma rine trans gres sion that started in early phases of the Badenian (i.e., Langhian). It tra di tion ally com prises a tri par tite di vi sion: pre-evaporitic, evaporitic (chem i - cal de pos its) and post-evaporitic strata (nu mer ous in for mal lithostratigraphic di vi sions that are dif fi cult to cor re late). The stra tig ra phy of this se quence is based mainly on plank tonic foraminifera and cal car e ous nannoplankton, which show that the bulk of the rocks may be cor re lated with the Mid dle Mio cene NN5 to NN9 cal car e ous nannoplankton zones and the M5 to M8–11 plank tonic foraminifera zones (e.g., Garecka et al., 1996; Garecka and Jugowiec, 1999; Olszewska, 1999).
A sim i lar, tri par tite di vi sion can be ap plied to the Mio cene suc ces sion in the Roztocze area, which rep re sents a mar ginal fa cies of the Carpathian Foredeep Ba sin. How ever, this suc - ces sion, due to its unique tec tonic po si tion, shows a dif fer ent
de vel op ment than its coun ter parts in cen tral ar eas of the sed i - men tary ba sin. Roztocze is an up lifted area (in re la tion to the Carpathian Foredeep to the south) in the south ern part of the Lublin Up land; its lon gi tu di nal struc ture stretches NW–SE from Kraœnik in Po land to Lviv in Ukraine (Fig. 1A). Due to its up lifted po si tion, the Mid dle Mio cene (Badenian) trans gres sion in Roztocze led to ac cu mu la tion of thin (merely 30–40 m thick com pared to over 2.000 m in the basinal part) shal low-ma rine coarse siliciclastic and organodetrital de pos its (Fig. 2). They rest upon Up per Cre ta ceous strata with lo cally pre served ma - rine Paleogene. Lo cally, in the base of the Mid dle Mio cene suc - ces sion, con ti nen tal phytogenic de pos its of un cer tain, pre sum - ably Lower Mio cene (Karpatian, i.e., up per most Burdigalian) age oc cur (e.g., Jankowski et al., 2014).
Both the ma rine Paleogene and con ti nen tal strata un der - went sig nif i cant ero sion dur ing and af ter up lift move ments re - spon si ble for for ma tion of the Roztocze struc ture. Ero sion led to al most com plete re moval of these strata. Most com plete sec - tions of the epicontinental ma rine Eocene in the Carpathian fore land are known from the So³okija Graben in Roztocze, and from the £ukowa-4 bore hole (Tarnogród vi cin ity) in the Fig. 1. Lo ca tion of the site stud ied and ge ol ogy of the study area
A – geo log i cal sketch of SE Po land show ing po si tions of main geo log i cal struc tures and lo cal i ties men tioned in the text (based on ¯ytko et al., 1989, from Oszczypko, 1996); B – lo ca tion of the ex po sure at Lipowiec (ar rowed)
Carpathian Foredeep. They rep re sent the Bartonian–Lower Priabonian in the So³okija Graben (GaŸdzicka, 1994; Gedl, 2014), and the Priabonian in the £ukowa-4 bore hole (Gedl, 2015). Iso lated sandy de pos its near Tomaszów Lubelski, which form a re sid ual hill, are be lieved to rep re sent the Paleogene, al - though no palaeontological ev i dence has been found so far (Buraczyñski and Krzowski, 1994).
The pres ence of Eocene ma rine strata at Roztocze and in the Carpathian Foredeep sug gests that the up lift of the Roztocze struc ture started in the Oligocene or later. Jankowski and Margielewski (2014) cor re lated phytogenic de pos its of Roztocze (Siemiatycze, Rawa Ruska) and the neigh bour ing part of the foredeep ba sin (Tarnogród), and con cluded that up - lift move ments be gan not ear lier than af ter their ac cu mu la tion in the Karpatian (Early Mio cene; this view re quires fur ther stud ies be cause at least some of phytogenic de pos its of the Carpathian Foredeep Ba sin may be of Oligocene age; Myœli wiec and Œmist, 2006; Gedl, 2015). Ac cord ing to Jankowski and Margie - lewski (2014), Roztocze was al ready an up lifted struc ture dur - ing de po si tion of organodetrital strata – dated to the Late Badenian ac cord ing to e.g., Szczechura (1982), Musia³ (1987), Wysocka (2002), and the Sarmatian–Pannonian ac cord ing to Czepiec (1996), and Jankowski and Margielewski (2015), who treated Badenian microfossils as re worked, al though no list of youn ger micro fossils was given by the lat ter.
MATERIAL
Ma te rial for this study co mes from sandy Mio cene strata that oc cur in a scarp of a steep top o graphic ridge of the Roztocze hills (Fig. 1B). Sam ples were col lected from an ex po - sure and a shal low bore hole lo cated a few metres north of the road that leads from the vil lage of Lipowiec to the top of a lo cal hill at the ham let of Lipowiec-Góry (GPS co or di nates:
N50°37.792’, E022°52.172’). This ex po sure, over 10 m high, was de scribed and il lus trated in de tail by Margielewski (in Kr¹piec et al., 2010: point 2) and Jankowski and Margielewski
(2014). It rep re sents basal part of the Badenian (Mid dle Mio - cene) transgressive suc ces sion at Roztocze (the Cre ta ceous sub strate was pen e trated by drill ing al most 9 m be low the bot - tom of the ex po sure; see Jankowski and Margie lewski, 2014:
fig. 4). The higher part of the sandy suc ces sion is ex posed some 100 m north, on the same side of the road; its top most part con sists of coralline al gal lime stone that forms the top of the hills.
Strata ex posed con sist of white, beige and rusty, non-cal - car e ous quartz sand that pre dom i nates in the up per part, and green ish non-cal car e ous quartz sand with glauconite in the basal part (Fig. 3). A few dark brown non-cal car e ous muddy clay lay ers oc cur. A hand-drilled bore hole pen e trated the un der - ly ing part of the suc ces sion, down to a depth of al most 5 m (i.e., it stopped ca. 4 m above the base of the Mio cene suc ces sion;
Fig. 3). This part con sists of dom i nat ing green ish, fine-grained non-cal car e ous glauconitic-quartz loamy sands interlayered with rusty coarse-grained quartz sand lay ers. A few pale green - ish clay lay ers are pres ent. A to tal of thir teen sam ples was taken from the ex po sure and core. Seven sam ples were taken from clay lay ers, three sam ples were taken from loamy sand with glauconite from the low er most drilled in ter val, and three sam ples from quartz sand from the up per part of the ex po sure (Fig. 3).
METHODS
The sam ples were pro cessed in the micropalaeontological lab o ra tory of the In sti tute of Geo log i cal Sci ences, Pol ish Acad - emy of Sci ences, Re search Cen tre in Kraków. The quan tity of rock pro cessed was vari able, de pend ing on the li thol ogy: 30 g for clay sam ples and 500 g for sand sam ples. Sam ples were washed in wa ter, and the frac tion be tween 10 and 250 µm (sieved at 250 and 10 µm on a ny lon mesh) was treated with 40% hy dro flu oric acid (HF), heavy-liq uid (ZnCl2 + HCl; den sity 2.0 g·cm–3) sep a ra tion. The res i due was sieved again at 10 µm on a ny lon mesh. No ni tric acid (HNO3) treat ment was ap plied.
Homotryblium-dominated Eocene dinoflagellate cyst assemblages from Middle Miocene (Badenian)... 463
Fig. 2. Syn thetic scheme of lithostratigraphy and li thol ogy of Mio cene strata in the Pol ish part of Roztocze (af ter Musia³, 1987, from Wysocka, 2002)
Two palynological slides were made from each sam ple us ing glyc er ine jelly as a mount ing me dium. All dinoflagellate cysts were counted from both slides us ing a Carl Zeiss Axiolab mi cro - scope. Pho to graphs of aquatic palynomorphs were taken us ing a Sony DSC-S75 cam era and Carl Zeiss Achroplan x100 oil lens. The rock sam ples, palynological res i dues and slides are stored in the col lec tion of the In sti tute of Geo log i cal Sci ences, Pol ish Acad emy of Sci ences, Re search Cen tre in Kraków.
RESULTS
Sam ples from quartz sand (5/2015) and quartz-glauconitic sand (3/2015) were bar ren. Loamy glauconitic sand (400, 470 and 490 cm) yielded or ganic de bris com posed of very fine par ti - cles of black and dark brown phytoclasts and pale re mains of land plant tis sues. Rare bisaccate pol len grains oc cur; no dinoflagellate cysts have been found.
Clay sam ples yielded higher amounts of palynological mat - ter, which, ex cept for sam ple 1/2015, in cludes aquatic palyno - morphs. The lat ter are ma rine dinoflagellate cysts pres ent in all sam ples, and thin-walled spher i cal forms of un cer tain or i gin found in the sam ple from 220 cm depth; their dis tri bu tion is shown in Fig ure 4. Dinoflagellate cysts are pale-col oured, their wall struc ture shows no traces of in creased ma tu rity. How ever, they are com monly pre served as frag ments (par tic u larly large chorate forms such as Areosphaeridium; Fig. 5P, X) or they show traces of me chan i cal dam age. The dinoflagellate cysts are il lus trated in Fig ures 5 to 7.
The sam ple from 450 cm depth yielded in fre quent frag - ments of un de ter min able dinoflagellate cysts, some of these pre sum ably be ing Cleistosphaeridium. The re main ing clay sam ples yielded as sem blages that con sist of fre quent Homo - tryblium rep re sent ing a mor pho log i cal com plex (H. floripes, H.
plectilum, H. vallum; see de Verteuil and Norris, 1996: p. 22;
Fig. 5A–O, Q–S) as so ci ated with var i ous pro por tions of other spe cies (Fig. 4). The pro por tion of Homotryblium in these sam - ples is as fol lows: 51% (380 cm), 96% (360 cm), 69% (220 cm), 88% (150 cm), 52% (2/2015), and 100% (4/2015).
Taxa other than the dom i nant Homotryblium are Glaphyro - cysta, Cleistosphaeridium, Heterosphaeridium, and Membra -
Fig. 3. Lipowiec ex po sure and bore hole with po si tion of sam ples col lected (ex po sure li thol ogy af ter Kr¹piec et al., 2010)
Fig. 4. Dinoflagellate cyst oc cur rence at Lipowiec
Homotryblium-dominated Eocene dinoflagellate cyst assemblages from Middle Miocene (Badenian)... 465
Fig. 5. Dinoflagellate cysts from sandy de pos its at Lipowiec (scale bar = 25 µm)
A–C – Homotryblium floripes (150 cm); D – Homotryblium plectilum (380 cm); E, F – Homotryblium floripes (150 cm); G, H – Homotryblium floripes (380 cm); I, J – Homotryblium plectilum (220 cm); K, L – Homotryblium vallum (220 cm); M–O – Homotryblium vallum (360 cm); P – Areosphaeridium diktyoplokum: iso lated paraplate with a pro cess ter mi nated with a char ac ter is tic plat form (2/2015); Q–S – Homotryblium vallum (360 cm); T – iso lated pro cess, pre sum ably of Areosphaeridium michoudii (380 cm); U, V – Areosphaeridium michoudii (380 cm); W – in com plete spec i men of Spiniferites pseudofurcatus (380 cm); X – iso lated pro cess, pre sum ably of Areosphaeridium michoudii (380 cm)
Fig. 6. Dinoflagellate cysts from sandy de pos its at Lipowiec (scale bar = 25 µm)
A, B – Glaphyrocysta sp. (380 cm); C, D – Glaphyrocysta semitecta (380 cm); E, F – Glaphyrocysta cf. semitecta (380 cm); G, H – Glaphyrocysta microfenestrata (sam ple 2/2015); I, J – Membranophoridium connectum (380 cm); K – Membranophoridium connectum (220 cm); L – Membranophoridium aspinatum (220 cm); M, N – Heterosphaeridium sp. A sensu Gedl, 2013 (380 cm); O, P – Heterosphaeridium sp. A sensu Gedl, 2013 (380 cm); Q–S – Heterosphaeridium sp. A sensu Gedl, 2013 (380 cm); T, X – Adnatosphaeridium multispinosum (220 cm); U–W – Adnatosphaeridium multispinosum (380 cm)
Homotryblium-dominated Eocene dinoflagellate cyst assemblages from Middle Miocene (Badenian)... 467
Fig. 7. Dinoflagellate cysts from sandy de pos its at Lipowiec (scale bar = 25 µm)
A–C – Cleistosphaeridium sp. A sensu Gedl, 2013 (380 cm); D – Cleistosphaeridium sp. A sensu Gedl, 2013: a wrin kled spec i men show ing gran u lar struc ture of the cyst wall (380 cm); E, F – Cleistosphaeridium cf. placacanthum (380 cm); G – Glaphyrocysta sp. (380 cm); H – Pentadinium? sp.: spec i men with thick and gran u lar endocyst (380 cm); I, J – Cleistosphaeridium cf. placacanthum sp. (380 cm); K, L – Homotryblium aculeatum (220 cm); M, N – Cleistosphaeridium placacanthum (150 cm); O, P – Cleistosphaeridium placacanthum (2/2015); Q–Y – spher i cal thin-walled palynomorphs (all 220 cm)
no phoridium, which are most fre quent in the sam ple from 380 cm depth. The sam ple from 220 cm depth is char ac ter ized by the fre quent oc cur rence of thin-walled prox i mate forms of un cer tain or i gin (Fig. 7Q–Y). Most of these are smooth, bi-lay - ered; none has a pylome. Some show a gir dle in equa to rial po - si tion, along which a split is ob served in some spec i mens (Fig.
7W). This split leads to a com plete sep a ra tion of two equal halves (Fig. 7X, Y). This fea ture re sem bles epicystal dino - flagellate cysts such as Mendicodinium, but in the case of the Lipowiec spec i mens no typ i cal dinoflagellate plate ar range ment (e.g., a sulcal notch) was ob served.
INTERPRETATION
BIOSTRATIGRAPHY
Mio cene strata of the Pol ish part of Roztocze have so far yielded rel a tively in fre quent dinoflagellate cysts. This is due to sed i men tary con di tions spe cific for this area, which in clude shal low-ma rine, high-en ergy con di tions (e.g., Wysocka, 2002) that were pre sum ably hos tile for dino flagellates and/or for their pres er va tion. Most of the coarse -grained, com monly organo - detrital rocks that build Roztocze are bar ren. Dinoflagellate cyst as sem blages are found in in fre quent fine-grained lay ers; they con sist there of a mix ture of re worked Eocene spe cies, and ones be lieved to be in situ. The sim i lar state of pres er va tion makes sep a rat ing these two groups dif fi cult in the case of long-rang ing spe cies. The as sem blages de scribed from Lipo - wiec con sist al most ex clu sively of spe cies that ap peared for the last time be fore the Mid dle Mio cene. A short over view of their ranges is given be low:
– Adnatosphaeridium multispinosum: Early–Late Eocene (Powell, 1992), Early–Mid dle Eocene (Stover et al., 1996), Eocene–Early Oligocene (Köthe and Piesker, 2007), Mid dle Eocene (Vasilyeva, 2013);
– Areosphaeridium diktyoplokum: lat est Early–Late Eo - cene (Powell, 1992), Early–Late Eocene (Stover et al., 1996; Köthe and Piesker, 2007); the last ap pear ance of this spe cies is re ported from the Eocene/Oligocene boun - d ary in ter val (see Eldrett et al., 2004, for dis cus sion);
– Areosphaeridium michoudii: Early Eocene (Vasilyeva, 2013); last ap pear ance: Mid dle Priabonian (Bujak and Mudge, 1994; Mudge and Bujak, 1994; Eldrett et al., 2004); Late Priabonian (Heil mann -Clausen and van Simaeys, 2005; Thom sen et al., 2012);
– Enneadocysta sp.: Mid dle Eocene–Early Oligocene (Sto ver et al., 1996 as the Enneadocysta arcuata “com - plex”; Köthe and Piesker, 2007 as Enneadocysta spp.);
– Glaphyrocysta microfenestrata: lat est Mid dle Eoce - ne–ear li est Oligocene (Powell, 1992), Early Oligocene:
Köthe and Piesker, 2007);
– Glaphyrocysta semitecta: Bartonian–Rupelian (De Co - ninck, 1995); Lutetian–Rupelian (Heilmann-Clausen and van Simaeys, 2005), Bartonian–Priabonian (Vasil y - eva, 2013); a to tal range of G. semitecta in North ern Hemi sphere mid-lat i tudes: lat est Lutetian–Early Rupe - lian (Wil liams et al., 2004); last ap pear ance: ear li est Rupelian (Mudge and Bujak, 1994);
– Membranophoridium aspinatum: Mid dle Eocene–Oligo - cene (Powell, 1992), Late Eocene–Early Oligocene (De Coninck, 1995), Late Eocene–Oligocene (Stover et al., 1996); Priabonian (Heilmann-Clausen and van Sima -
eys, 2005); Munsterman and Brinkhuis (2004) re port the high est oc cur rence of this spe cies from up per most Chattian–Lower Aquitanian of the North Sea.
A Paleogene, most likely Eocene age can be re ferred to the spe cies Cleistosphaeridium sp. A and Heterosphaeridium sp.
A, both de scribed by the au thor from the Eocene Popiele Beds (Gedl, 2013).
Pre cise dat ing of the dinoflagellate cyst as sem blages in ques tion is en abled by Homotryblium floripes, and the mor pho - log i cally sim i lar H. plectilum and H. vallum, which are most fre - quent in the ma te rial stud ied. Be ing widely dis trib uted in Paleo - gene and Early Mio cene strata, the oc cur rence of Homotry blium oc cur rence in the Mid dle and Up per Mio cene is less ev i dent.
Dybkj³r and Piasecki (2010) re ported an acme of Homo tryblium (in clud ing H. plectilum and H. tenuispinosum) from the up per - most Oligocene–low er most Mio cene of the North Sea. A sim i lar acme of Homotryblium floripes/plectilum was re ported by Munsterman and Brinkhuis (2004) from the co eval in ter val of the North Sea Mio cene. The same au thors re ported the high est oc - cur rence of Homotryblium vallum in the Lower Mio cene of the North Sea (up per most Aquitanian–low er most Burdigalian).
A sim i lar age of this event is given by de Verteuil and Norris (1996). Stover et al. (1996) show a Lower Mio cene high est oc - cur rence of H. plectilum and H. vallum in north west ern Eu rope and off shore east ern North Amer ica. Powell (1986) re ports a Chattian last ap pear ance of H. plectilum and Aquitanian last ap - pear ance of H. floripes in NW It aly. Ac cord ing to Wil liams et al.
(1993) both H. floripes and H. vallum ap peared for the last time in North At lan tic in late Early Mio cene. Sev eral au thors (e.g., Wrenn and Kokinos, 1986; Dybkj³r and Piasecki, 2010) re - ported the oc cur rence of Homotryblium in the Up per Mio cene or even youn ger strata, but ac cord ing to de Verteuil and Norris (1996: p. 20, 22; see this pub li ca tion for full ref er ence list and dis - cus sion) these re cords are pre sum ably the re sult of re work ing.
First ap pear ance data of the Homotryblium floripes com plex show that these spe cies ap peared for the first time in the Eocene, mainly dur ing the Late Eocene. Wil liams et al. (2004) re port a to tal range of H. floripes from equa to rial ar eas as Lutetian–Late Mio cene, but in higher lat i tudes its old est oc cur - rences are known from the Priabonian (Heilmann-Clausen and van Simaeys, 2005; Vasilyeva, 2013) and Bartonian–?Pria - bonian (Gedl, 2014, 2015). H. vallum, in turn, is known al ready from Eocene strata, e.g., up per most Priabonian (Köthe and Piesker, 2007), and Bartonian–Priabonian (Gedl, 2013).
The dis tri bu tion of Homotryblium in the Mio cene of the Carpathian Foredeep Ba sin sup ports the the sis of de Verteuil and Norris (1996). This ge nus is miss ing in most Mid dle Mio - cene (Badenian) sec tions stud ied, and if pres ent, it oc curs as ev i dently re worked (poorly pre served from the Carpathian Paleo gene; e.g., Gedl, 1999) or as so ci ated with other ev i dently re worked spec i mens (e.g., Gedl, 1999, 2005, 2016). Soli man and Piller (2007), who stud ied the Karpatian/Badenian bound - ary in ter val in the Styrian Ba sin in Aus tria, noted the rare oc cur - rence of Homotryblium sp. in Karpatian strata only; Badenian strata, de void of re worked Paleogene forms, con tained no Homotryblium.
The spe cies with the lon gest strati graphic ranges among those found in the sam ples stud ied are Cleistosphaeridium placacanthum – Mid dle Eocene to Late Mio cene (e.g., Eaton et al., 2001) and Spiniferites pseudofurcatus – Cre ta ceous–Mid - dle Mio cene (e.g., Stover et al., 1996). Both spe cies could thus be in situ. They have been en coun tered as in situ forms in the Up per Badenian of Pol ish Roztocze, where they co-oc cur with Eocene spe cies (Gedl, 2016). They are also known from co eval strata of the Ukrai nian part of the Carpathian Foredeep Ba sin,
where they oc cur in as sem blages de void of re worked spec i - mens (Gedl and Peryt, 2011) or they are as so ci ated with rare re worked Paleogene spec i mens (Peryt et al., 2014).
This over view of strati graphic ranges sug gests that dino - flagellate cyst as sem blages from Lipowiec are of Late Eocene age. The ques tion of whether C. placacanthum and S. pseudo - furcatus are in situ or they are re worked re mains un solved; the lack of any other Mio cene spe cies, like those from youn ger strata of Roztocze (e.g., Polysphaeridium), sug gests that they are also re worked.
PALAEOENVIRONMENT
The lack of Mio cene dinoflagellate cysts in the de pos its stud ied points to a re stricted sed i men tary set ting. Too lit tle data is avail able for a pre cise en vi ron men tal re con struc tion, but the most likely fac tor hos tile for dinoflagellates and/or their pres er - va tion was the very shal low wa ter and high-en ergy hy dro dy - namic con di tions. The lat ter, eas ily vis i ble in sed i men tary struc - tures (e.g., Jankowski and Margielewski, 2014), may be re - spon si ble for the lack of dinoflagellate cysts. Dur ing pe ri ods of calmer de po si tion, clay lay ers ac cu mu lated. They yielded re - worked spec i mens only; the lack of in situ taxa shows that en vi - ron men tal con di tions were still not fa vour able for dino fla - gellates. The pres ence of incertae sedis forms (spher i cal paly - no morphs) in the sam ple from 220 cm depth, pos si bly in situ, may be re lated to these spe cific sed i men tary con di tions, but so long as their palaeo environmental pref er ences are un known, noth ing more can be sug gested.
Re worked dinoflagellate cysts can be much more use ful, rep re sent ing the only palaeontological trace of eroded Eocene strata. Al though their as sem blages may be in com plete due to se lec tive ero sion and/or re-ac cu mu la tion pro cesses, on their ba sis an at tempt of Late Eocene Roztocze en vi ron ment re con - struc tion can be undertaken.
The pres ence of ma rine as sem blages shows that this part of Rozotcze was also flooded by a ma rine trans gres sion dur ing the Late Eocene. The lack of off shore spe cies such as Impa - gidinium (e.g., Wall et al., 1977; Harland, 1983; Ed wards and Andrle, 1992; Brinkhuis, 1994; Rochon et al., 1999; Boessen - kool et al., 2001) and Nematosphaeropsis (Brinkhuis, 1994;
Dale, 1996; Rochon et al., 1999) sug gests a near-shore, prox i - mal ma rine set ting. This in ter pre ta tion is sup ported by fre quent oc cur rences of Homotryblium. This ge nus is widely as so ci ated with near-shore, lit to ral en vi ron ments, com monly with in crea - sed sa lin ity (e.g., Köthe, 1990; Brinkhuis, 1994). The lat ter fea - ture may in di cate slightly in creased sa lin ity of the ma rine wa - ters, al though Dybkj³r (2004) sug gested that Homotryblium may also ben e fit from de creased sa lin ity con di tions.
An other char ac ter is tic fea ture of the Lipowiec as sem blages is a lack of peridinioids, which are mainly heterotrophs and thrive in wa ters with in creased nu tri ent avail abil ity (e.g., deltaic or upwelling wa ters; see e.g., Biffi and Grignani, 1983; Brin - khuis, 1994; Sprangers et al., 2004; Sluijs et al., 2005). Their lack may thus re flect oligotrophic con di tions. This re con struc - tion is sup ported by a lack of Lingulodinium machaero phorum, an autotroph, which is com monly as so ci ated with eutrophic prox i mal wa ters (e.g., Targarona et al., 1999; Boessen kool et al., 2001).
All these fea tures sug gest that dinoflagellate cysts found in sands at Lipowiec in hab ited a near shore, pre sum ably shal low ma rine set ting with rather oligotrophic wa ters with slightly in - creased sa lin ity. There fore, a la goonal embayment can be re - con structed for this part of Roztocze dur ing the Late Eocene.
DISCUSSION
This is the first re port of dinoflagellate cyst oc cur rence in the basal part of the Mid dle Mio cene (Badenian) suc ces sion in this part of Roztocze. At tempts to search for dinoflagellate cysts in co eval de pos its un der taken by the au thor dur ing the last few years have proved fruit less: quartz sands ex posed at Gleboviti and Stradc (Ukrai nian part of Roztocze; see Wysocka and Roniewicz, 2004: fig. 3 for cor re la tion) were found to be bar ren.
Data pre sented in this and pre vi ous pa pers show that dur ing the Badenian trans gres sion in the Pol ish part of Roztocze in tense ero sion of ma rine Eocene strata took place. Clay lay ers within Up per Badenian organodetrital lime stone in the vi cin ity of Józefów yielded fre quent Mid dle and Late Eocene dino flagellate cysts (Gedl, 2012). Jankowski et al. (2014) re ported cal car e ous nannoplankton typ i cal of the NP 17 Zone (Barto nian) from clays over ly ing Mio cene brown-coal de pos its in Siedliska.
The Lipowiec dinoflagellate cyst as sem blages show qual i ta - tive and quan ti ta tive sim i lar i ties to the re worked as sem blages from Józefów in the fre quent oc cur rence of Homotryblium floripes com plex rep re sen ta tives (Gedl, 2016). Three sam ples col lected from the Józefów quar ries yielded the fol low ing pro - por tions of H. floripes plus H. plectilum and H. vallum (cal cu - lated for re worked forms only): 49% (sam ple Pds-1), 55%
(Pds-2), and 30% (Jzf-3). High pro por tions of Homotryblium and a si mul ta neous lack of peridinioids sug gest that dur ing both the Early and Late Badenian sim i lar la goonal Eocene set tings were grad u ally washed out in this part of Roztocze. The lower pro por tion of Homotryblium in the Up per Badenian sam ples, as so ci ated with a higher num ber of taxa such as Areo - sphaeridium michoudii, Areoligera spp., and Batiacasphaera
?compta (gen. et spec. indet. sensu Gedl, 2013), may in di cate a grad ual ero sion of the Eocene suc ces sion in Roztocze, and their sub se quent in verted ac cu mu la tion dur ing the Badenian.
Dur ing the Early Badenian the top of the Priabonian suc ces sion un der went ero sion (and hence the youn gest as sem blages com posed al most en tirely of Homotryblium were washed out and ac cu mu lated); dur ing the Late Badenian, in turn, ero sion reached the base of the Eocene suc ces sion (Barto - nian–?Priabonian), and older as sem blages with less com mon Homotryblium were eroded. Ac cep tance of this sce nario sug - gests that the orig i nal Eocene suc ces sion in this part of Roztocze ac cu mu lated in a grad u ally shallowing ma rine en vi - ron ment, grad u ally trans form ing to a la goonal embay ment, re - flected by the in creas ing pro por tion of Homotryblium.
Re worked dinoflagellate cysts from Lipowiec show some tax o nomic sim i lar i ties (par tic u larly when Homotryblium floripes com plex rep re sen ta tives are con sid ered) to Eocene as sem - blages that oc cur in situ at Roztocze. An as sem blage with com - mon Homotryblium floripes, dated to the lat est Barto - nian?–Early Priabonian, was de scribed by the au thor from the top of the Eocene suc ces sion of the So³okija Graben in Roz - tocze; on the base of its com po si tion, a very shal low ma rine, pre sum ably partly re stricted set ting was re con structed (Gedl, 2014). The Homotryblium floripes-dom i nated So³okija as sem - blages dif fer, how ever, by the co-oc cur rence of peridinioids and Lingulodinium machaerophorum, which are miss ing in ma te rial from Lipowiec. More over, strata from the top of the So³okija, which con tain these as sem blages, are sep a rated by lay ers that yielded al most monospecific Deflandrea as sem blages.
The oc cur rence of peridinioids is also typ i cal of the Priabonian suc ces sion in the Carpathian Foredeep (£ukowa-4 bore hole); rare spec i mens of Lingulodinium machaerophorum have been found there (Gedl, 2015). Al though these strata Homotryblium-dominated Eocene dinoflagellate cyst assemblages from Middle Miocene (Badenian)... 469
yielded fre quent spec i mens of Homotryblium too, in this case they rep re sent H. aculeatum; H. floripes is rare there.
Homotryblium plectilum, in turn, is a com mon spe cies in Priabonian dark mudstones of the Popiele Beds at Koniusza (Gedl, 2013). The sed i men tary set ting of these mudstones was in ter preted as a mar ginal, prox i mal, near-shore zone of the Carpathian sea. Also in this case, peridinioids (mainly Deflan - drea), al though sub or di nate, oc cur as an im por tant part of dinoflagellate cyst as sem blages (Gedl, 2013: figs. 13 and 14).
The com par i son pre sented above shows that the Lipowiec as sem blages are no ta ble for the lack of peridinioids and Lingulodinium machaerophorum. The other as sem blages from co eval strata, al though also in clud ing fre quent Homotryblium, con tain peridinioids and L. machaerophorum. Oc cur rences of the two lat ter spe cies pre sum ably re flect slightly higher avail - abil ity of nu tri ents, pos si bly caused by a higher in flux of fresh - wa ter. This is clearly vis i ble, par tic u larly in case of the So³okija suc ces sion, where lev els with Homotryblium are interlayered with lev els with monospecific Deflandrea as sem blages, likely re flect ing brack ish con di tions.
The as sem blages from Lipowiec are thus pre sum ably fur - ther traces of Late Eocene near-shore ma rine set tings that sur - rounded the emerged ar eas to the north of the pres ent day Lublin Up land; to the south they passed into the deep ma rine bas ins of the Carpathian sea. These set tings were char ac ter - ized by fre quent palaeoenvironmental changes, typ i cal of shal - low-ma rine, prox i mal ma rine en vi ron ments, where la goonal, tidal, deltaic and other sedimentological set tings com monly re - placed or interfingered with each other. The Lipowiec as sem - blages pre sum ably re flect la goonal oligotrophic en vi ron ments, whereas co eval as sem blages from neigh bour ing ar eas in hab - ited settings influenced by freshwater influx.
CONCLUSIONS
1. The sandy suc ces sion that oc curs in basal parts of the Mio cene at Roztocze usu ally con tains no fos sils and its strati - graphic po si tion is based on su per po si tion (Musia³, 1987). The re sults pre sented in this pa per are the first doc u men ta tion of their dinoflagellate cyst con tent. Their as sem blages, how ever, are reworked.
2. Lack of Mio cene dinoflagellate cysts in de pos its stud ied sug gests that en vi ron men tal con di tions dur ing early stages of the Badenian trans gres sion at Roztocze were un fa vour able for dinoflagellates and/or in ap pro pri ate for their pres er va tion (e.g., high en er getic hy dro dy namic con di tions dur ing coarse grain fraction accumulation).
3. The dis cov ery of re worked Eocene dinoflagellate cysts in the Lower Badenian, the basal part of suc ces sion, as de scribed in this pa per, shows that ero sion of the ma rine Paleogene cover started si mul ta neously with the Badenian trans gres sion.
4. The tax o nomic com po si tion of the re worked as sem - blages shows that they were washed out from near-shore, shal - low-wa ter Up per Eocene de pos its ac cu mu lated pre sum ably in a la goonal embayment char ac ter ized by oligotrophic high-sa lin - ity wa ters. Ero sion of sim i lar de pos its, al though pre sum ably slightly older, lasted also dur ing the Late Badenian when
organo detrital car bon ate de pos its ac cu mu lated in this part of Roztocze.
5. Com par i son of the Lipowiec as sem blages with Late Eocene dinoflagellate cyst as sem blages from other sites at Roztocze and the Carpathian fore land shows that the lat ter, al - though also shal low-ma rine and partly la goonal, ac cu mu lated in slightly dif fer ent en vi ron ments. Lack of taxa that bene fited from high con cen tra tion of nu tri ents in ma rine wa ters, such as peridinioids and Lingulodinium machaerophorum, sug gest oligotrophic con di tions in the case of Lipowiec. The oc cur rence of both peridinioids and L. machaerophorum (in var i ous pro por - tions) at other sites shows that these ar eas were in flu enced, to a cer tain de gree, by fresh wa ter in flux in the Late Eocene.
Ac knowl edge ments. I would like to thank A. Wysocka (Uni ver sity of War saw) for tell ing me about the ex po sure of glauco nitic sand at Lipowiec. W. Margielewski (Pol ish Acad emy of Sci ences, Kraków) is ac knowl edged for valu able com ments and dis cus sion while pre par ing this manu script. B. S³odkowska (Pol ish Geo log i cal In sti tute – Na tional Re search In sti tute, Warszawa), S. De Sche pper (Uni ver sity of Bergen) and an anon y mous re viewer are kindly ac knowl edged for their crit i cal re marks on the manu script.
SPECIES LIST
(taxonomy and citations follow Fensome et al., 2008):
Adnatosphaeridium multispinosum Wil liams et Downie, 1966
Areosphaeridium diktyoplokum (Klumpp, 1953) Eaton, 1971
?Areosphaeridium michoudii Bujak, 1994
Cleistosphaeridium placacanthum (Deflandre et Cook - son, 1955) Eaton et al., 2001
Cleistosphaeridium cf. placacanthum (Deflandre et Cook son, 1955) Eaton et al., 2001
Cleistosphaeridium sp. A sensu Gedl, 2013 Enneadocysta sp.
Glaphyrocysta microfenestrata (Bujak, 1976) Stover et Evitt, 1978
Glaphyrocysta semitecta (Bujak, 1980) Lentin et Wil - liams, 1981
Glaphyrocysta cf. semitecta (Bujak, 1980) Lentin et Wil - liams, 1981
Glaphyrocysta sp.
Heterosphaeridium sp. A sensu Gedl, 2013 Homotryblium aculeatum Wil liams, 1978
Homotryblium floripes (Deflandre et Cookson, 1955) Stover, 1975
Homotryblium plectilum Drugg et Loeblich Jr., 1967 Homotryblium vallum Stover, 1977
Membranophoridium aspinatum Gerlach, 1961 Membranophoridium connectum Stover et Hardenbol, 1994
Pentadinium? sp.
Spiniferites pseudofurcatus (Klumpp, 1953) Sarjeant, 1970
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