Geo log i cal Quar terly, 2017, 61 (4): 887–895 DOI: http://dx.doi.org/10.7306/gq.1379
Late Albian cal car e ous dinocysts and calcitarchs re cord linked to en vi ron men tal changes dur ing the fi nal phase of OAE 1d – a case study
from the Tatra Moun tains, Cen tral West ern Carpathians
Agnieszka CIUREJ1, Krzysztof BĄK1, * and Marta BĄK2
1 Ped a gog i cal Uni ver sity of Cra cow, In sti tute of Ge og ra phy, Podchorążych 2, 30-084 Kraków, Po land
2 AGH Uni ver sity of Sci ence and Tech nol ogy, Fac ulty of Ge ol ogy, Geo phys ics and En vi ron men tal Pro tec tion, Al. Mickiewicza 30, 30-059 Kraków, Po land
Ciurej, A., Bąk, K., Bąk, M., 2017. Late Albian cal car e ous dinocysts and calcitarchs re cord linked to en vi ron men tal changes dur ing the fi nal phase of OAE 1d – a case study from the Tatra Moun tains, Cen tral West ern Carpathians. Geo log i cal Quar - terly, 61 (4): 887–895, doi: 10.7306/gq.1379
Cal car e ous dinocysts and calcitarchs have been in ves ti gated for the first time within the Up per Albian lime stone and marl suc ces sion of the Zabijak For ma tion from the High-Tatric Unit in the Tatra Moun tains (Cen tral West ern Carpathians), re lated to the Oce anic Anoxic Event 1d (OAE 1d). Four groups of morphotaxa of cal car e ous dinocysts have been dis tin guished.
They to tally dom i nate the as sem blages, and be long to the pithonellids. They are rep re sented by Pithonella sphaerica (Kaufmann in Heer) and P. ovalis (Kaufmann in Heer), which dom i nate, as well as P. trejoi Bonet and P. lamellata Keupp in Keupp and Kienel, which are less abun dant. Two other morphotaxa, Colomisphaera gigantea (Borza) and Cadosina oraviensis Borza, oc cur spo rad i cally in the as sem blages. Both forms rep re sent the calcitarch group, which as sem bled calcispheres of un known tax o nomic af fin ity. The cal car e ous dinocyst and calcitarch di ver sity is low to mod er ate, com pared to the gen eral spe cies rich ness known from Late Albian as sem blages in other West ern Tethyan sec tions. This is in ter preted as a re sult of nu tri ent in put fluc tu a tions due to changes in the cir cu la tion pat tern of sur face and in ter me di ate wa ters. The changes in the P. sphaerica/P. ovalis ra tio along the Up per Albian sec tion are here cor re lated with short-term (third-or der) sea level fluc tu a tions in clud ing transgressive and re gres sive events and a highstand. Pelletization pro cesses might have in - flu enced cyst abun dance on the sea floor, es pe cially dur ing pe ri ods with oligotrophic sur face wa ters.
Key words: cal car e ous dinocysts, calcitarchs, nu tri ent in put and sea level fluc tu a tions, Up per Albian, OAE 1d, Tatra Moun - tains.
INTRODUCTION
The Late Albian was a time of en hanced ma rine burial of sed i men tary or ganic car bon as re lated to in or ganic car bon, which is char ac ter ized by a pos i tive ex cur sion in d13C val ues in ma rine de pos its, known from var i ous Tethyan, North At lan tic, Pa cific and epicontinental en vi ron ments, and at trib uted to Oce - anic Anoxic Event 1d (e.g., Coccioni et al., 2001; Strasser et al, 2001; Ken nedy et al., 2004; Bornemann et al., 2005; Jarvis et al., 2006; Gale et al., 2011; Papp et al., 2013; Scott et al., 2013;
Horikx et al., 2014; Melinte-Dobrinescu et al., 2015). Such strata (a lime stone and marl suc ces sion) with a pos i tive d13C ex cur sion, de pos ited on the north ern shelf of the Al pine–Carpa - thian microcontinent, were also re corded from the Cen tral West ern Carpathians (the Tatra Moun tains; Bąk et al., 2016).
They con tain ma rine and land-de rived or ganic mat ter, strongly
de graded un der oxic con di tions. These car bon ate de pos its, Late Albian in age, are en riched in cal car e ous dinocysts. How - ever, there are rel a tively few pub li ca tions which deal with cal - car e ous dinocysts from the Cre ta ceous Tethyan realm of the Carpathians, with the ex cep tion of Up per Ju ras sic through Lower Cre ta ceous as sem blages (mostly Tithonian–Aptian) re - corded in nu mer ous West ern Carpathian set tings (e.g., Nowak, 1963, 1968, 1974; Borza, 1964, 1969, 1986; Řehánek, 1987;
Reháková and Michalík, 1994, 1996; Lintnerová et al., 1997;
Michalík et al., 1999, 2009; 2016; Reháková, 2000a, b;
Pszczółko wski and Myczyński, 2004; Olszewska, 2005; Olsze - wska et al., 2008; Pszczółkowski et al., 2016).
In this pa per, we pro vide the first over view of cal car e ous dinocysts and calcitarchs, to tally dom i nated by pithonellids from the Up per Albian car bon ate de pos its of the High-Tatric Unit in the Cen tral West ern Carpathians. Us ing new palaeontological data re lated to these microfossils from the Tatra Moun tains, we make sug ges tions as to the pos si ble en vi ron men tal con di tions pre vail ing on the north ern shelf of the Al pine – Carpathian microcontinent (Fig. 1) that in flu enced the de vel op ment of these or gan isms dur ing the time cor re spond ing to Oce anic Anoxic Event 1d.
* Corresponding author, e-mail: sgbak@cyf-kr.edu.pl Received: February 13, 2017; accepted: July 29, 2017; first published online: September 11, 2017
CALCAREOUS DINOFLAGELLATES AND CALCITARCHS
Sin gle-celled spher i cal to ovoid, cal car e ous microfossils known in the fos sil re cord from the Pa leo zoic to the Ce no zoic have been usu ally re ferred to as “calcispheres” (Wil liam son, 1880). Dur ing over a cen tury of palaeontological stud ies, many microfossils pre vi ously as signed to “calcispheres” have been clas si fied to var i ous tax o nomic groups (see Kaźmierczak and Kremer, 2005). They were con sid ered to be long to var i ous groups of foraminifers, cal car e ous al gal spores, pro to zo ans, spores of dasyclad al gae or nannoplankton. Many forms have been proved to rep re sent dinoflagellate cal car e ous cysts as rest ing, re pro duc tive or coccoid stages of their life cy cle (Meier et al., 2009) be long ing to the Fam ily Thoracosphaeraceae (Elbrächter et al., 2008). De spite many pre vi ous in ves ti ga tions, the bi o log i cal af fin i ties of many “calcispheres” re main un known.
For these forms Versteegh et al. (2009) pro posed a new incertae sedis group of the Calcitarcha that in cludes all cal car e - ous microfossils with a cen tral cav ity and cur rently lack ing tax o - nomic al lo ca tion.
Pithonellids, which are the dom i nant cal car e ous micro - fossils in the as sem blage stud ied, have been con sid ered to be - long to cal car e ous dinoflagellates (Elbrächter et al., 2008;
Wendler et al., 2013; Wendler and Bown, 2013). Two other forms – Colomisphaera gigantea (Borza) and Cadosina ora - viensis Borza – which are pres ent in the ma te rial stud ied, rep re - sent the calcitarch group in which are as sem bled “calcispheres”
of un known tax o nomic af fin ity.
GEOLOGICAL SETTING
The Cen tral West ern Carpathians are com posed of three prin ci pal superunits: the Tatric, Veporic, and Gemeric base - ment cover crustal im bri cates (Plašienka, 1999). The Tatric superunit (Tatricum) frames the outer (north ern) part of the Cen tral West ern Carpathians block (microcontinent), formed dom i nantly by pre-Al pine crys tal line base ment com plexes and a Late Pa leo zoic–Me so zoic (up to Lower Turonian) sed i men - tary cover. The de pos its stud ied be long to the High-Tatric units
oc cur ring in the Tatra Moun tains which are a part of the Tatric superunit (Plašienka, 2003). The High-Tatric units are ex posed in three tec tonic-fa cies units: Kominy Tylkowe, Czerwone Wier - chy and Giewont (e.g., Kotański, 1961). The Kominy Tylkowe Unit (called the autochthonous unit) rests di rectly on the crys tal - line base ment, and com prises a Lower Tri as sic (?Perm ian) through mid-Cre ta ceous sed i men tary suc ces sion (e.g., Rabo - wski, 1959; Kotański, 1961; Lefeld, 1968; Jurewicz, 2005;
Jezier ska et al., 2016, Wolska et al., 2016). Its youn gest part, the Zabijak For ma tion (Krajewski, 2003) con tains echino - dermal -foraminiferal lime stones of the Żeleźniak Mem ber, which are over lain by marls and marly lime stones of the Kamienne Mem ber. The Żeleźniak Mem ber and the low er most part of the Kamienne Mem ber, both late Albian in age (Bąk and Bąk, 2013) con tain abun dant cal car e ous dinocysts which are an ob ject of this study.
MATERIAL AND METHODS
The sec tion stud ied is lo cated in the Żeleźniak gully, a left trib u tary of the Kościeliska Val ley in the Pol ish part of the Tatra Moun tains (Fig. 2). The ex po sure is sit u ated at the bot tom of the gully, and on its slopes (Fig. 2C), ex tend ing for a dis tance of 100 m be low a 3 m high wa ter fall. The lower part of the sec tion com prises the up per most part of a lime stone suc ces sion (Żeleźniak Mem ber), ~30 cm thick. This is over lain by grey and dark grey marls fol lowed by green marls (Kamienne Mem ber).
We stud ied cal car e ous dinocysts and calcitarchs in thin - -sec tions 5 x 3 cm in size made from rock sam ples. Four rock sam ples rep re sent a 30 cm thick echinodermal-foraminiferal lime stone layer, and 13 sam ples rep re sent a 4 m marly in ter val that com prises 2 m of dark grey marls fol lowed by 2 m of green marls (Fig. 3). The rock ma te rial from the same strati graphi cal po si tion has been a sub ject of ear lier stud ies, which con cerned bio- and chemostratigraphy, as well as or ganic mat ter anal y ses (Bąk, 2015; Bąk et al., 2015; Bąk et al., 2016).
Con tents of plank tonic foraminifers, pel lets and phos phates are shown as per cent val ues of the to tal microfacies com po - nents. The to tal con tent of pithonellids and calcitarchs was es ti - mated on the ba sis of reg u lar tra verses at 200 mm in ter vals across a slide area of ~6 cm2. Tax o nomic stud ies were based on ob ser va tions of their mor pho log i cal fea tures such as shape, size, ap er ture, and wall struc ture (mostly, the wall crys tal ori en - ta tion) fol low ing the de ter mi na tions known from lit er a ture (e.g., Keupp, 1987, 1990; Reháková, 2000a, b; Wendler et al., 2002a, 2013; Kohring et al., 2005; Wendler and Bown, 2013;
OmaÔa et al., 2014).
STRATIGRAPHY
The plank tonic foraminiferal as sem blages stud ied in the Żeleźniak Gully sec tion and the nearby sec tions (Bąk and Bąk, 2013; Bąk, 2015) show that the whole in ter val cor re sponds to the lower part of the Parathalmanninella appenninica Zone (Up - per Albian; Fig. 3; for dis cus sion – see Bąk et al., 2016). The pos i tive ex cur sion in d13C val ues re corded in the lime stone layer and first marly in ter val (Bąk et al., 2016), with val ues com - pa ra ble to that seen in the GSSP sec tion at Mont Risou, east of Rosans, Hautes-Alpes (Ken nedy et al., 2004; Gale et al., 2011), and Speaton, York shire, Eng land (Mitch ell et al.,1996), in di cate that this part of the sec tion cor re sponds to Oce anic Anoxic Event 1d.
888 Agnieszka Ciurej, Krzysztof Bąk and Marta Bąk
Fig. 1. Up per Albian (~100 Ma) palaeogeographic map sho w - ing the lo ca tion of the strata stud ied (red as ter isk) in the West ern Tethys (map re drawn af ter http://deeptime - maps.com – sim pli fied); names of re gions partly af ter Micha - lik et al., 2007)
Alp.-Carp. Or. – Al pine–Carpathian Orogen;
Vep. L. – Veporic Land
RESULTS
Calcispheres ap pear in all sam ples across the sec tion stud - ied (Fig. 3). Their to tal con tent ranges from 5 up to 20% of microfacies con stit u ents in lime stone and grey marl cor re - spond ing to the OAE 1d in ter val. Above this in ter val, in green marlstones, their con tent di min ishes and fluc tu ates be tween 3 and 7%. A char ac ter is tic fea ture of these de pos its is the oc cur - rence of plank tonic foraminifers, dom i nated by hedbergellids and less com mon heterohelicids (Fig. 4A, B), as well as of two types of pel let – ho mo ge neous and not di gested, in clud ing nearly com plete tests of plank tonic foraminifers and pithonellids (Fig. 4C, D).
Six morphotaxa of “calcispheres” were rec og nized within the sec tion stud ied (Figs. 3 and 5). The as sem blages are dom i - nated by cal car e ous dinocysts, which are rep re sented by the ge nus Pithonella Lorenz, 1902 (Fig. 5) be long ing to P. ovalis (Kaufmann in Heer), and P. sphaerica (Kaufmann in Heer), which com prise >95% of the in di vid u als ob served. Less abun - dant are P. trejoi Bonet and P. lamellata Keupp in Keupp and Kienel. There are also forms which are here clas si fied as Colomisphaera gigantea (Borza) and Cadosina oraviensis Borza be long ing to calcitarchs but dis play ing the pithonellid wall spec trum. The first one in cludes a large forms cur rently ques - tion ably clas si fied to the Cil i ates. The spec i mens of Cadosina oraviensis Borza con tain forms with walls that ap pear to be dom i nated by an or ganic sub stance. The pre cise tax o nomic po si tion of these morphotaxa is im pos si ble to de ter mine at this state of study due to the low num ber of in di vid u als.
The spe cies rich ness of the calcitarch and cal car e ous dinocyst as sem blages ranges from 3 to 6 in an in di vid ual sam - ple (Figs. 3 and 6). They are gen er ally mod er ately- to well-pre - served even if the rock ma trix was recrystallized (Fig. 5).
Based on the changes in as sem blage dis tri bu tion, three in - ter vals can be dis crim i nated in the sec tion in ves ti gated. In ter val 1 and In ter val 2 cor re spond to the OAE 1d event. In ter val 1 com prises light grey echinoderm-foraminiferal lime stone con -
tain ing nu mer ous pel lets and phos phates (Fig. 6). “Calcis - phere” as sem blages are rep re sented only by pithonellids rang - ing from 2 to 4 spe cies per sam ple. Pithonella ovalis pre vails over other taxa. Ly ing above, In ter val 2 con tains grey marls in - clud ing abun dant pel lets (Fig. 6) and land-de rived flo ral com po - nents (Bąk et al., 2016). Cal car e ous dinocysts and a calcitarch as sem blage is rich in this in ter val con sist ing of up to 55% of all con stit u ents (sam ple Z8) and highly di verse (spe cies di ver sity up to 6; Fig. 6); Pithonella sphaerica with thick, dou ble-lay ered, cal car e ous walls dom i nates here con sist ing of nearly 70% of the as sem blages. In ter val 3 is rep re sented by green marls ly ing above the OAE 1d de pos its. It is again char ac ter ized by the dom i nance of P. ovalis with an over all lower spe cies num ber, up to 3 (Fig. 6).
The P. sphaerica/P. ovalis ra tio that is used as a bathymetric in di ca tor in ma rine en vi ron ments (e.g., Dias-Brito, 2000; Wendler et al., 2002b) fluc tu ates along the sec tion stud - ied (Fig. 6) rang ing from 0.2 to 2.7 (av er age value per in ter vals, de scribed above). The high est val ues cor re sponds to the dark grey marls of the OAE 1d in ter val.
The fluc tu a tions in pithonellid dis tri bu tion cor re spond also to fluc tu a tions in the to tal num ber of pel lets vis i ble in thin-sec tion views, es pe cially in those sam ples which con tain whole or partly bro ken plank tonic foraminiferal tests. How ever, this dis tri bu tion does not co in cide with to tal cal cium car bon ate con tent (Bąk et al., 2016; Fig. 6).
DISCUSSION
The “calcispheres” from the sec tion stud ied are dom i nated by the pithonellid group (>95% of the whole as sem blages). The in ter pre ta tion of palaeoenvironment based on pithonellid ecol - ogy match ob ser va tions by pre vi ous au thors that these photosynthetic or gan isms lived in sur face wa ters and are as so - ci ated with sed i ments de pos ited in the shelf to shal low bathyal en vi ron ments (e.g., Ban ner, 1972; Zügel, 1994; Wendler et al., Late Albian cal car e ous dinocysts and calcitarchs re cord linked to en vi ron men tal changes dur ing the fi nal phase of OAE 1d... 889
Fig. 2A – Carpathians against the back ground of a sim pli fied geo log i cal map of the Al pine orog eny and their fore land;
B – lo ca tion of the area stud ied in the Tatra Moun tains (In ner Carpathians) on a con tour map (map af ter Bryndal, 2014);
C – lo ca tion of the sec tion stud ied in the Żeleźniak Gorge
2002a, b; OmaÔa et al., 2014; Wiese et al., 2015). There are state ments that Pithonella ovalis is a char ac ter is tic spe cies of the off shore fa cies and marks transgressive in ter vals. In turn, Pithonella sphaerica com monly oc curs in an en vi ron ment that is slightly in flu enced by coastal con di tions (e.g., Zügel, 1994).
Dur ing the Late Cenomanian–Early Turonian, both these taxa showed sig nif i cant and tem po ral blooms, known as the global bioevent as so ci ated with OAE 2 (e.g., Wendler et al., 2002a, b). The abun dance of pithonellids was strongly re lated to nu tri ent avail abil ity in sur face wa ters. Jarvis et al. (1988) stated that this group ap peared to have been an op por tu nis tic group and their abun dance in sed i ments prob a bly re flects an in - creased nu tri ent sup ply; more over, the marked in crease of them co in cides with a de cline in or ganic-walled dinoflagellate cysts. Al ter na tive sug ges tions have been pre sented by
Dias-Brito (2000) and Wiese et al. (2015), who con cluded that the pithonellids are typ i cal of dis tal, most likely nu tri ent-de - pleted, shelf en vi ron ments. In turn, OmaÔa et al. (2014) de - scrib ing the large quan tity of forms from the ge nus Pithonella in the Cenomanian–Turonian de pos its of car bon ate plat form from Mex ico sug gested that these microfossils were op por tun ists liv - ing in an un sta ble en vi ron ment. An other palaeo eco logi cal as - pect con cern ing these microfossils was sug gested by Keupp (1987). He re marked a change of “calcispheres to wards pitho - nellid dom i nated as sem blages dur ing the Cenomanian trans - gres sion. Sim i larly, Wendler et al. (2002a, b) used the ra tio of P.
sphaerica to P. ovalis (Ps/Po) as an in di ca tor in the in ter pre ta - tion of rel a tive dis tances from the coast line. They pos tu lated that a high Ps/Po ra tio (~10) char ac ter izes mar ginal shelf as - sem blages, while a low Ps/Po ra tio (~3), may sug gest that the as sem blages came from the outer shelf.
CALCAREOUS DINOCYSTS VS. SEA LEVEL CHANGES AND NUTRIENT AVAILABILITY
Tak ing into ac count the Ps/Po ra tio in three in ter vals dis tin - guished along the sec tion stud ied, we in ter pret that the pitho - nellids rep re sent the outer shelf as sem blages sensu Wendler et al. (2002a, b; av er age Ps/Po ra tio ranges be tween 0.2 and 2.7) that was also ear lier sug gested based on the com po si tion of ben thic foraminiferal as sem blages (B¹k, 2015).
The oc cur rence of pithonellids dur ing the pe riod re lated to In ter val 1 cor re sponds to a transgressive phase that cul mi nated in the highstand event. It is sup ported in the Tatric area by fa - cies rep re sented by the echinoderm-foraminiferal lime stone, rich in its top most part in phos phates with hardgrounds, stromatolites, and phos phate pisolites (sum mary in Krajewski, 2003). This short-term (third-or der) transgressive event be gan dur ing the P. ticinensis Chron (af ter KAl7 us ing a num ber ing sys tem of Cre ta ceous eustatic events af ter Haq, 2014; Fig. 6) and cul mi nated dur ing the P. appenninica Chron. The up per wa ter col umn was char ac ter ized by high pro duc tiv ity in sur face wa ters, po ten tially in duced by wind-driven coastal upwelling (B¹k et al., 2016). In this en vi ron ment, in put of un di gested pel - lets to the sea bot tom de rived di rectly from su per fi cial wa ters was mod er ate, which cor re lates with the mod er ate spe cies di - ver sity of cal car e ous dinocysts (Fig. 6). Such a mod er ate value of “calcisphere” di ver sity is prob a bly di min ished due to long - -last ing ma te rial re work ing and dis so lu tion on the sea floor, caused by a long pe riod of sed i ment ex po sure and nondepo - sition be fore the diagenesis of the sed i ment (B¹k, 2011).
The pithonellids from In ter val 2 dis play the high est val ues of the Ps/Po ra tio in the sec tion stud ied (av. value ranges from 1.4 to 2.7; Fig. 6). This co in cides with the high abun dance of all types of “calcispheres”, and ad di tion ally, is linked with the abun - dance of plank tonic foraminiferal tests be long ing mostly to the hedbergellids and heterohelicids (Figs. 4A, B and 6). The lat ter microfossils were typ i cal com po nents of mid-Cre ta ceous su - per fi cial oce anic wa ter oc cur ring abun dantly dur ing blooms of
“calcispheres”, re lated to transgressive events (e.g., Keupp, 1987; OmaÔa et al., 2014). How ever, tak ing into ac count the change in the sed i men ta tion re gime dur ing this pe riod, re - corded in the com po si tion of the dark grey marl suc ces sion that over lies the echinoderm-foraminiferal lime stone, it seems that the blooms of “calcispheres” were linked rather to a high land-de rived nu tri ent de liv ery in su per fi cial wa ter. These de pos - its con tain nu mer ous clay min er als and two types of or ganic mat ter (OM), in clud ing ma rine with abun dant non-di gested or partly di gested pel lets, and land-de rived, with com po nents of near-shore al lu vial plains in the sur round ing land scape, i.e.,
890 Agnieszka Ciurej, Krzysztof B¹k and Marta B¹k
Fig. 3. Lithological log of the Up per Albian suc ces sion from the ¯eleŸniak Gully sec tion, High Tatric units,
Tatra Moun tains, plot ted against the dis tri bu tion of pithonellids and calcitarchs
ZM – ¯eleŸniak Mem ber
vas cu lar plant rem nants, spores of lake-de rived green al gae and par ti cles of fresh wa ter blue al gae (for de tails – see Bąk et al., 2016). Nu tri ents de rived from rivers may have stim u lated the growth of phytoplankton and, thus, zoo plank ton ac tiv ity and en hanced the de po si tion of POC (fe cal pel lets and ag gre - gates). The oc cur rence of autochthonous bar ite crys tals and ho mo ge neous pel lets in these de pos its may in di cate rel a tively high pro duc tiv ity in the sur face wa ters. In turn, the change in the wa ter re gime from weak en ing coastal upwelling to downwelling dur ing that time caused wa ter strat i fi ca tion that is in ter preted here based on the en hanced con tent of un di gested pel lets in the de pos its along the In ter val 2 (Fig. 6). The flux of such pel lets into the bot tom sed i ments might have been a con se quence of sec ond ary di ges tion by microzooplankton in the wa ter col umn be low the thermocline/nutricline, and ad di tion ally, by mi cro bial de com po si tion at the sea bot tom (Pomeroy et al., 1984).
Fluc tu a tions in the to tal num ber and di ver sity of pithonellids and calcitarchs in In ter val 2 is an other fea ture of these as sem - blages. This may re cord changes in sur face wa ter cur rents re - lated to changes in wind di rec tion, caus ing vari able sea sonal an oxia and wa ter-col umn strat i fi ca tion. The high est num ber of highly di verse “calcispheres” that co in cides with the re duced per cent age of plank tonic foraminifers oc cur ring in the dark grey marls with or ganic-rich laminae con tain ing land-de rived OM (sam ples Z4b and Z8; Fig. 6) is an ex am ple of con di tions with a less ox y gen ated sea floor, caused by co pi ous in put of OM from
al lu vial plains cov ered by wetlands. It seems, there fore, that the nu tri ents brought by rivers to the sea may have the great est in - flu ence on the growth of pithonellids and calcitarchs.
It should be stressed that the source of echinoderm par ti - cles, which were nu mer ous in the un der ly ing suc ces sion of the lime stones, dis ap peared at the be gin ning of In ter val 2 due to burial by land-de rived clay ma te rial, trans ported to gether with land-de rived or ganic de tri tus. This shows that the growth of
“calcispheres”, de scribed from In ter val 2, oc curred dur ing the continuing highstand event.
The abun dance and di ver sity of pithonellids and calcitarchs as sem blage di min ished dur ing In ter val 3. This co in cides with the low av er age value of the Ps/Po ra tio (0.6–1.2) and the re - duced con tent of un di gested pel lets in the green marlstones (Fig. 6). Most prob a bly, the “calcispheres” from these sed i - ments rep re sent the as sem blages that cor re spond to the pe riod dur ing which the wa ter strat i fi ca tion was bro ken down and the upwelling re gime was re newed. This has been con firmed by many geo chem i cal in di ces based on anal y sis of OM in the green marlstones, which show a de crease in the sup ply of land-de rived OM to the ba sin floor, and ad di tion ally sug gest more highly ox y gen ated con di tions on the ba sin floor (Bąk et al., 2016). All this may in di cate that the weaker growth of
“calcispheres” dur ing this pe riod cor re sponds to the be gin ning of a third-or der re gres sion event, still with en hanced nu tri ent avail abil ity in sur face wa ter.
Late Albian cal car e ous dinocysts and calcitarchs re cord linked to en vi ron men tal changes dur ing the fi nal phase of OAE 1d... 891
Fig. 4. Char ac ter is tic microfacies in the Up per Albian marl suc ces sion from the Żeleźniak Gully sec tion, High Tatric Unit, Tatra Moun tains
A, B – pho to mi cro graphs of foraminiferal-pithonellid microfacies in dark grey marlstone from the up per most part of the OAE 1d in ter val; Hb – Hedbergella, Ht – Heterohelix; sam ple Z8; C, D – pel lets pro duced by micro - zooplankton with vis i ble in trin sic con stit u ents, which are plank tonic foraminifers and pithonellids, C – sam ple Z6, D – sam ple Z11
892 Agnieszka Ciurej, Krzysztof Bąk and Marta Bąk
Fig. 5. Cal car e ous dinoflagellate cysts and calcitarchs from the Up per Albian de pos its in the Żeleźniak Gully sec tion, High Tatric units, Tatra Moun tains
A–D – Pithonella lamellata Keupp in Keupp and Kienel, A – sam ple Z1d; B – sam ple Z1h; C – sam ple Z2b; D – sam ple Z4b; E, F – Pithonella ovalis (Kaufmann in Heer) – lon gi tu di nal sec tion show ing the eliptical shape, thick wall and small ap er ture at the end (ar row), E – sam ple Z3b, F – sam ple Z6; G–J – Pithonella sphaerica (Kaufmann in Heer), G – ax ial sec tion, sam ple Z2b; H – sam ple Z4b; I – sam ple Z4b; J – ax ial sec tion show ing the dou ble layer of the thick wall di vided by a dark line (black ar row) and ap er ture (white ar row), sam ple Z5;
K – Pithonella trejoi Bonet – lon gi tu di nal sec tion with ap er ture (ar rows), Z1b; L – Colomisphaera gigantea (Borza) with ap er ture (ar row) – a large form of the pithonellid wall spec trum, sam ple Z12; M – Cadosina oraviensis Borza – form with pithonellid wall spec trum, rich in an or ganic sub stance, ar row points to the ap er ture, sam ple Z2b
CONCLUSIONS
The “calcispheres” re corded from the Up per Albian de pos - its of the Tatra Moun tains are of rel a tively low to mod er ate di - ver sity, and con sist of four spe cies of pithonellids and two spe - cies be long ing to the calcitarchs. The as sem blages are dom i - nated by pithonellids, the rel a tive abun dance of which ex ceeds 95%. The dis tri bu tion of “calcispheres” in the sec tion stud ied can be re lated to changes in nu tri ent in put that oc curred pe ri od - i cally, be ing re lated to changes in sea level dur ing the Late Albian. The al ter nat ing abun dance of P. sphaerica and P. ovalis gives a prox i mal/dis tal proxy that shows short-term third-or der fluc tu a tions of sea level dur ing that time, which took place be - tween two mid-Cre ta ceous sea level fall events, de scribed as KAl7 and KAl8 fol low ing the chronostratigraphy by Snedden and Liu (2011). A dom i nance of P. ovalis is char ac ter is tic for both transgressive and re gres sive in ter vals, in di cat ing a gen - eral outer shelf po si tion. Pe ri ods of mod er ate abun dances of
“calcispheres” can be in ter preted to re flect in creas ing mar ginal upwelling near the north ern slopes of the sub merged Tatric Ridge, where sur face and nu tri ent-rich wa ters were mixed. Dur - ing the highstand pe riod, wa ter strat i fi ca tion was con du cive to oligotrophy, cre at ing the most fa vour able con di tions for the in - creased de po si tion of pithonellid cysts.
Ac knowl edge ments. The study was sup ported by the Min - is try of Sci ence and Higher Ed u ca tion to A. Ciurej (Pro ject DS-UP-WGB No. 4n), by the Na tional Sci ence Cen tre to K. B¹k (grant 2011/01/B/ST10/07405), and by the Min is try of Sci ence and Higher Ed u ca tion to M. B¹k (Pro ject DS-AGH Uni ver sity of Sci ence and Tech nol ogy, WGGiOŒ-KGOiG no.
11.11.140.005). The au thors are grate ful to J. Wendler, D. Reháková and an anon y mous re viewer for their help ful re - marks. Spe cial thanks go to K. Leszczyñski for im prov ing the Eng lish text.
Late Albian cal car e ous dinocysts and calcitarchs re cord linked to en vi ron men tal changes dur ing the fi nal phase of OAE 1d... 893
Fig. 6. Lithological log of the Up per Albian suc ces sion from the ¯eleŸniak Gully sec tion, High Tatric Unit, Tatra Moun tains, plot ted against se lected geo chem i cal and micropalaeontological in di ces
d13Ccarb, CaCO3, phos phates and pel lets con tents – af ter B¹k et al. (2016); homog. – ho mo ge neous; n/d – un di gested;
OAE 1d – Oce anic Anoxic Event 1d – grey back ground; for ex pla na tions of lith o logic log – see Fig ure 3
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