Downloaded from geology.gsapubs.org on January 7, 2014 Long-term ecosystem stability in an Early estuary

Martin Zuschin1*, Mathias Harzhauser2, Babette Hengst1, Oleg Mandic2, and Reinhard Roetzel3 1University of , Department of Palaeontology, Althanstrasse 14, A-1090 Vienna, 2Natural History Museum Vienna, Department of Geology and Palaeontology, Burgring 7, A-1010 Vienna, Austria 3Geological Survey of Austria, Neulinggasse 38, A-1030 Vienna, Austria

ABSTRACT southern estuarine part, probably connected to The question of ecosystem stability is central to ecology and paleoecology and is of particu- a huge, south-north–trending river system in the lar importance for estuaries, which are environmentally highly variable, considered as geo- adjoining Vienna Basin, and a northern marine logically short lived, and among the most degraded modern ecosystems of our planet. Under- part (Harzhauser et al., 2002; Latal et al., 2006). standing their ecological dynamics over geological time scales requires paleontological data in A connection to the large epicontinental Parate- a sequence stratigraphic framework, which allows evaluation of paleocommunity dynamics thys Sea was most probably established along in an environmental context. A 445-m-thick estuarine succession in a satellite basin of the the northeastern margin of the Korneuburg Ba- Vienna Basin (Austria) shows continuous sedimentation over 700 k.y. and can be divided into sin (Fig. 1) (Harzhauser and Wessely, 2003). The two transgressive systems tracts and a highstand systems tract. In contrast to expectations, foraminifera in parts of the basin were adapted no major physical disturbances of the ecosystem involving abrupt changes in diversity and to brackish-water conditions and indicate a very biofacies composition occurred at fl ooding surfaces and at the sequence boundary. Accom- shallow water environment through basin his- modation space remained remarkably constant over the depositional history of the basin, tory, with maximum water depth not exceeding and all changes between depositional environments were therefore more or less gradational. 30 m (Rögl, 1998). The fi sh fauna (e.g., Gobi- Biotic change along the studied succession can be described as a gradual faunal replacement idae, Sparidae, Dasyatidae, Myliobatidae) in the in response to habitat tracking, a process also reported for some normal marine shelf environ- southern Korneuburg Basin indicate littoral and ments. Benthic assemblages in the estuarine succession were strongly dominated by a few taxa shallow sublittoral conditions in a subtropical to and developed along two indirect gradients, water depth and hydrodynamic energy. These tropical environment with freshwater infl uence gradients show subtle long-term trends, corresponding to the sequence stratigraphic architec- (Reichenbacher, 1998). ture. Tectonics affected the sequence architecture in this particular marginal marine setting: it controlled accommodation space and sedimentary input, and provided stable boundary STRATIGRAPHY conditions over hundreds of thousands of years. Our study demonstrates for the fi rst time The section near the village of Stetten in the that estuaries, which are under great environmental pressure today, are resilient to natural Korneuburg Basin can be divided into Stetten environmental perturbations and can persist over geological time scales. West (total thickness of 445 m) and Stetten East (total thickness of 165 m) (Fig. 1; Fig. DR1 in INTRODUCTION and belong to the most degraded modern eco- the GSA Data Repository1). Thin coaly depos- The fossil record is rich in examples of eco- systems of our planet (Lotze et al., 2006). Be- its, washed-in land-, and freshwater snails in- system stability exceeding millions of years, cause of their highly variable physicochemical dicate marginal marine conditions. Sand pack- mostly in Paleozoic and Mesozoic marine characteristics, estuaries also count as naturally ages with trough cross-bedded sets are mostly level bottom communities, interrupted by brief stressed areas: their biotas have the ability to interpreted as tidal sand waves of the shoreface. episodes of strong turnover (DiMichele et al., adapt to various stressors and the ecosystem can Pelitic sediments mostly show even lamination 2004). Ecosystem stability at the scale of tens compensate for changes in the environment, a to wavy bedding or thinly alternating sandy and to hundreds of thousands of years is a typical feature termed homeostasis (Elliott and Quinti- muddy layers, indicative of tidal fl at deposits feature of Pleistocene coral reefs (Pandolfi and no, 2007). The study of turnover and communi- (Reineck and Singh, 1975; Boyd et al., 2006). Jackson, 2006), but has only rarely been report- ty stability in such a fl uctuating ecosystem over The succession is dominated by upward-fi ning ed from Phanerozoic soft bottom assemblages hundreds of thousands of years is therefore of parasequences, which are typical for a tidal fl at (Holterhoff, 1996). High-resolution studies of major ecological and paleoecological interest. to subtidal environment on a muddy, tide-domi- this kind require a stratigraphic framework to nated shoreline (Van Wagoner et al., 1990). The evaluate paleocommunity dynamics in their en- STUDY AREA astronomical tuning of the gamma ray record vironmental context (Holland and Patzkowsky, The Korneuburg Basin, a satellite basin of the fi xed the deposition to the time interval from 2004; Scarponi and Kowalewski, 2004). Here Vienna Basin, is located in and is 17.0 to 16.3 Ma, yielding a total duration of we report for the fi rst time on ecosystem stabil- 20 km long and 7 km wide (Fig. 1). It was formed 700 k.y. (our data). ity over 700 k.y. in a Miocene estuarine succes- by Burdigalian pull-apart movements within the sion (Fig. 1). Alpine-Carpathian thrust belt. The basin sedi- SAMPLE PREPARATION Estuaries are semienclosed coastal water ments mostly belong to the Miocene Korneuburg We took a total of 118 bulk samples, weigh- bodies with salinities that differ from the open Formation, which was dated into nannoplankton ing ~1 kg each, from 96 shell beds. The sedi- sea. They exhibit distinct biota (Whitfi eld and zone NN4, paleomagnetic chron C5C, and mam- ment material >1 mm mesh size was quanti- Elliott, 2011), are environmentally highly vari- mal zone MN5 (Harzhauser and Wessely, 2003). tatively picked under a binocular microscope able (Elliott and Quintino, 2007), are considered Faunal composition and stable isotopes show for all biogenic components. For sponges, ev- as geologically rather short lived (Wolff, 1983), that the Korneuburg Basin was divided into a ery bioeroded biogenic hard part with distinct

*E-mail: [email protected]. 1GSA Data Repository item 2014002, Table DR1 (abundance and species richness of higher taxa), Table DR2 (environmental and stratigraphic affi liation of samples in biofacies), Figure DR1 (section Stetten East), Figures DR2–DR4 (results of two-way cluster analysis and ordination [nMDS]), and Appendix DR1 (actualistic comparison of abundant taxa), is available online at www.geosociety.org/pubs/ft2014.htm, or on request from [email protected] or Documents Secretary, GSA, P.O. Box 9140, Boulder, CO 80301, USA.

GEOLOGY, January 2014; v. 42; no. 1; p. 7–10; Data Repository item 2014002 | doi:10.1130/G34761.1 | Published online 6 December 2013 ©GEOLOGY 2013 Geological | January Society 2014 of America.| www.gsapubs.org Gold Open Access: This paper is published under the terms of the CC-BY license. 7 Downloaded from geology.gsapubs.org on January 7, 2014

STETTEN WEST Sequence Rarefied Natural radioactivity Lithology stratigraphy Biofacies DC 1 DC 2 species richness (gamma log) 123 12 4 6 8 10 16° 20' Lithology 440 )2 m depth energy Sand TS shallow high low high Silt-Clay T(

Vienna Interbedding Kleinebers- 48° 30' 2 dorf N Sand-Silt Lignite c t AUSTRIAAUSTRIA WE

Flysch basement a

S 400 r

t

-400 m

Sequence Stratigraphy s

m

-100 m

Roseldorf Zone e

parasequences t

retrogradational s

parasequence sets y

360 s aschberg Unit Swell aggradational to W progradational

e

rg Basin parasequence sets v 0 m u

i i

s

s

Schliefberg fault

e e

KorneubKorneuburg Basin r Stetten fault 320 g 123 12 4 6 8 10

-600 m

286-324 m s

Stetten Profile gap n a a 123 12 4 6 8 10

ienna Basin r

0 m V Korneuburg 48° 280 Flysch Unit 20' T Rhenodanubian 5 km

240 270 Samples Biofacies Intertidal 240 Agapilia biofacies Granulolabium- 210 Agapilia biofacies high energy Subtidal 180 Nassarius-Turritella- 200 Corbula biofacies 150 Nassarius-Paphia- Loripes biofacies DCA 2 DCA

120 ec

ssa ne

90 rr ucco 160 m

60 of land snails land of H i g h s t a n d y e m r c (HST) 30

) low energy

1 0

T

0 40 80 120 160 200 240 280 320 S

T( 1 1 T( deep DCA 1 shallow 120

t t

c c

Taxa Cliona designating taxa

400 a of intertidal biofacies t r Porites

Megabalanus designating taxa 320 Paphia of subtidal biofacies e m s Donax Peronaea Balanus Cerithium Lasaeina Ocenebra 240 Timoclea AcanthocardiaCrassostrea Sandbergeria 80 t Perna Terebralia

Loripes Sparidae Hydrobia

s

Granulolabium s y 160 Polinices Perrona Circomphalus Pyrene Nassarius Cubitostrea Turritella Cingula

DCA 2 DCA 80 Nucula C.arcellum e

Striarca Pelecyora Anadara Agapilia v Natica Antalis C.praeplicata s i 0 Cyllenina

Acteon 40

s s

Lesueurigobius Clavatula -80 e Bittium r Corbula Turboella

g s s g -160 Parvicardium

n a r r a n -160 -80 0 80 160 240 320 400 480

DCA 1 0 T 123 12 4 6 8 10 50 30 10 cps Figure 1. Korneuburg Basin, Austria, with studied transect and results of detrended correspondence analysis (DCA). Upper left: Sediment thickness and major structural units in region of basin (after Wessely, 1998). Arrows indicate freshwater infl ow in south and connection with open sea in north. Center left: Ordination plot of DCA of samples, showing four biofacies. Bottom left: Ordination plot of DCA of taxa. Right panel: Transect of Stetten West, including sequence stratigraphy, distribution of biofacies, axis 1 and 2 scores from DCA, species richness, and gamma log (cps—counts per second). Transect consists of two fourth-order sequences. Lower transgressive systems tract TST1 shows succession from tidal fl at to shoreface conditions from base to top; highstand systems tract consists of intertidal and very shallow subtidal pelites and sands, including root horizons and coal seams; upper TST2 is similar to TST1, but basal 30 m and top were not exposed. Major coal deposit is interpreted as sequence boundary (SB). Maximum fl ooding surface in both TSTs consists of package of sand deposits, interpreted as sand waves transported by tidal currents. For much shorter and more proximal section Stetten East, see Figure DR1 (see footnote 1).

8 www.gsapubs.org | January 2014 | GEOLOGY Downloaded from geology.gsapubs.org on January 7, 2014 traces of the ichnogenus Entobia was counted typically inhabit subtidal environments, and 4.0 A B as a sponge colony. Each balanid plate, decapod those with higher scores inhabit the intertidal or claw, echinoid spine, tooth, and otolith, and for very shallow subtidal environments (Appendix 2.8 2 1 AB 3 NTCB molluscs every shell (gastropods, scaphopods) DR1 in the Data Repository). The complex indi- 2 GAB 4 NPLB 1.6 TST2 and every isolated valve (bivalves), was counted rect gradient water depth is therefore the domi- 4 as an individual. In addition to these quantitative nant source of faunal variation in this study and 3 1 Log abundance HST TST1 samples, each layer was scoured for large-sized most likely refl ects the combined infl uence of 0.4 species, which are likely underrepresented in many environmental parameters that are highly 10 30 50 70 90 110 24 48 72 96 bulk samples, and a species was added as pres- correlated (Patzkowsky and Holland, 2012, p. Rank Rank ent to the data matrix where appropriate. Our 56). Sediment grain size is a poor predictor for C NTCB D approach biases relative importance toward taxa the composition of benthic assemblages (R = 100 TST1 NPLB TST2 AB with disarticulated body parts. The patterns of 0.12, p = 0.001; Fig. DR3) and DC2 is therefore 60 HST GAB our study, however, are consistent when correc- best interpreted as hydrodynamic energy, where 20

tions for disarticulation are applied. Abundance taxa with high scores (>100) prefer higher water No. of species 5000 15000 5000 15000 and species richness are strongly dominated by energy and fi rm or hard substrata, and taxa with Number of shells Number of shells molluscs (Table DR1 in the Data Repository). low scores (<100) prefer lower water energy and Figure 2. Rank abundance distribution (RAD), Each sample was classifi ed as either pelitic or soft substrata. Accordingly, low DC1 and DC2 rarefaction curves (RC), and species rich- sandy and assigned to a sequence and systems sample scores represent quiet water and mostly ness. A: RAD of systems tracts (HST—high- tract (ST) (Table DR2). pelitic subtidal environments, which are char- stand systems tract; TST—transgressive acterized by the Nassarius-Turritella-Corbula systems tract). B: RAD of biofacies. C: RC of STs. D: RC of biofacies. AB—Agapilia bio- STATISTICAL METHODS biofacies (NCTB). High DC1 and low DC2 val- facies; GAB—Granulolabium-Agapilia biofa- Species contributing <0.1% to the total as- ues represent quiet water environments of an cies; NCTB—Nassarius-Turritella-Corbula semblage, samples with <20 individuals, and inner tidal fl at, characterized by the Agapilia bio facies, NPLB—Nassarius-Paphia-Loripes one sample consisting solely of washed-in river biofacies (AB). Low DC1 and high DC2 scores, biofacies. snails were removed from the fi nal data matrix in turn, represent relatively high energy environ- (108 samples, 45 species, and 97.6% of the orig- ments of the shoreface, mostly characterized inal data). To standardize samples, percentages by sandy substrates and the Nassarius-Paphia- much higher than that of the HST and the GAB of the abundances were calculated and square Loripes biofacies (NPLB). High DC1 and DC2 and the AB. root transformed to de-emphasize the infl uence sample scores represent somewhat higher ener- of the most abundant taxa. getic conditions of the outer tidal fl at, character- DISCUSSION The benthic assemblages were explored for ized by the Granulolabium-Agapilia biofacies Biofacies changes are usually gradational differences between sediment composition and (GAB) (Fig. 1; Fig. DR3). within parasequences, but abrupt across fl ood- STs using analysis of similarity (ANOSIM). All four biofacies occur in both sections. The ing surfaces, typically either within TSTs or Because of the large sample size, differences be- AB mostly occurs in the highstand systems tract when a sequence boundary is associated with a tween sample groups were signifi cant, although (HST) and the GAB mostly in transgressive sys- major shift in facies (Holland, 2000). The ordi- low R-values indicate only poor discrimination. tems tract 1 (TST1) of the lower sequence. A nation in this study (Fig. 1), however, indicates Results of ANOSIM are virtually identical when few samples of the GAB were also present in no disjunct change in biofacies composition corrections for disarticulation are applied. Two- the HST of the lower sequence, but it was not within the TSTs or across the SB. Our interpre- way cluster analysis (Ward’s method) was ap- present in TST2 of the upper sequence. This is tation is that accommodation space remained plied to characterize biofacies and to detect hier- probably because the lower (i.e., shallower) part remarkably constant over the depositional his- archical groupings within the data set; detrended of this ST, where this biofacies would be ex- tory of the basin. Water depth varied only within correspondence analysis (DCA) and nonmetric pected to occur, is missing. The NCTB is almost the narrow limits of few tens of meters, and all multidimensional scaling were used as comple- exclusively present in the two TSTs. Similarly, changes between depositional environments mentary ordination methods to detect ecological the NPLB is almost exclusively restricted to the are therefore more or less gradational. Accord- gradients (Patzkowsky and Holland, 2012, p. two TSTs (Fig. 1; Table DR2). ingly, biodiversity also changed only little over 66). Cluster analysis and ANOSIM are based on Benthic assemblages are barely separable be- the studied succession, and most changes can the Bray-Curtis similarity coeffi cient. Logarith- tween STs (R = 0.189, p = 0.001; Fig. DR3), and be related to the presence or absence of envi- mic scale rank abundance plots and rarefaction DCA axes scores and diversity of samples show ronments within STs. The biotic change along curves were used to compare community orga- only weak trends across the studied sections. the studied succession is termed faunal replace- nization between the total assemblages of STs DC1 scores decrease from bottom to top of ment, a process that can only be observed in and biofacies. Mean rarefi ed species richness of TST1 in both sections. Apart from some distinct continuous sections. Such replacement typically samples (n = 20, according to smallest sample outliers, they are uniformly high in the HST occurs over thousands to hundreds of thousands size in the data set) was used to track diversi- and decrease again in TST2. Along the transect of years and most likely refl ects habitat track- ties within and between STs. Statistical analy- DC2 scores are similarly high in the two TSTs, ing, where species migrate to preferred habi- ses were performed with the software packages but typically lower in the HST. Rarefi ed spe- tats along environmental gradients (Brett et al., PRIMER version 6.1.6 (Clarke and Warwick, cies richness of samples is more variable in the 2007). Ecosystem stability here is exemplifi ed 1994) and PAST (Hammer et al., 2001). two TSTs and often above 8; it is less variable by recurrence of biofacies in STs. The strong and, with few exceptions, below 7 in the HST dominance of a few taxa in all STs and biofacies RESULTS (Fig. 1; Fig. DR1). STs and biofacies all show indicates that the benthic habitats covered in our Four overlapping biofacies are developed strong dominance of a few taxa, especially in study are physically controlled. This is typical along two major environmental gradients (Fig. 1; the GAB and the AB and in TST1 and the HST for estuaries, where the environment fl uctu- Figs. DR2 and DR3). DC1 is interpreted as wa- (Fig. 2). Evenness of the total assemblage of the ates constantly (Elliott and Quintino, 2007). As ter depth, because taxa with low scores (<130) two TSTs and of the NCTB and the NPLB is can be expected in such settings, evenness is

GEOLOGY | January 2014 | www.gsapubs.org 9 Downloaded from geology.gsapubs.org on January 7, 2014 higher in the two subtidal biofacies and in the fl uctuations: A review of faunal replacement and Miocene, Korneuburg Basin, Austria): Interna- two TSTs. the process of habitat tracking: Palaios, v. 22, tional Journal of Earth Sciences, v. 95, p. 95– During the studied interval the strong tectonic p. 228–244, doi:10.2110/palo.2005.p05-028r. 106, doi:10.1007/s00531-005-0510-3. Clarke, K.R., and Warwick, R.M., 1994, Change in Lotze, H.K., Lenihan, H.S., Bourque, B.J., Bradbury, reorganization of the central Paratethys resulted marine communities: An approach to statisti- R.H., Cooke, R.G., Kay, M.C., Kidwell, S.M., in rapid subsidence of the Korneuburg Basin cal analysis and interpretation: Plymouth, UK, Kirby, M.X., Peterson, C.H., and Jackson, J.B.C., (Wessely, 1998). Tectonics therefore affected Plymouth Marine Laboratory, 144 p. 2006, Depletion, degradation, and recovery poten- sequence architecture in this particular mar- DiMichele, W.A., Behrensmeyer, A.K., Olszewski, tial of estuaries and coastal seas: Science, v. 312, T.D., Labandeira, C.C., Pandolfi , J.M., Wing, p. 1806–1809, doi:10.1126/science.1128035. ginal marine setting by controlling subsidence S.L., and Bobe, R., 2004, Long-term stasis in Pandolfi , J.M., and Jackson, J.B.C., 2006, Eco- and sedimentary input, and by providing stable ecological assemblages: Evidence from the logical persistence interrupted in Caribbean boundary conditions over 700 k.y. fossil record: Annual Review of Ecology and coral reefs: Ecology Letters, v. 9, p. 818–826, Systematics, v. 35, p. 285–322, doi:10.1146/ doi:10.1111/j.1461-0248.2006.00933.x. CONCLUSIONS annurev.ecolsys.35.120202.110110. Patzkowsky, M.E., and Holland, S.M., 2012, Strati- Elliott, M., and Quintino, V., 2007, The estuarine graphic paleobiology: Understanding the distri- Ecosystem stability over geological time quality paradox, environmental homeostasis bution of fossil taxa in time and space: Chicago, scales is shown here for the fi rst time for envi- and the diffi culty of detecting anthropogenic London, University of Chicago Press, 259 p. ronmentally highly variable estuaries. We dem- stress in naturally stressed areas: Marine Pollu- Reichenbacher, B., 1998, Fisch-Otolithen aus dem onstrate that under appropriate tectonic bound- tion Bulletin, v. 54, p. 640–645, doi:10.1016/j Karpat des Korneuburger Beckens: Beiträge zur .marpolbul.2007.02.003. Paläontologie, v. 23, p. 325–345. ary conditions, these ecosystems, which are Hammer, O., Harper, D.A.T., and Ryan, P.D., 2001, Reineck, H.-E., and Singh, I.B., 1975, Depositional sed- strongly affected by anthropogenic impact to- PAST: Paleontological Statistics Software Pack- imentary environments: Berlin, Springer, 439 p. day and considered as geologically short lived, age for Education and Data Analysis: Palae- Rögl, F., 1998, Foraminifernfauna aus dem Karpat can persist over hundreds of thousands of years, ontologia Electronica, v. 4, p. 9 (http://palaeo (Unter-Miozän) des Korneuburger Beckens: a pattern previously only reported for some -electronica.org/2001_1/past/issue1_01.htm). Beiträge zur Paläontologie, v. 23, p. 325–345. Harzhauser, M., and Wessely, G., 2003, The Karpa- Scarponi, D., and Kowalewski, M., 2004, Stratigraphic normal marine level bottom assemblages of the tian of the Korneuburg Basin (Lower Austria), paleoecology: Bathymetric signature and se- shelf and coral reefs. The stability of such a low- in Brzobohatý, R., et al., eds., The Karpatian— quence overprint of mollusk associations from diversity ecosystem over such a long time and A lower Miocene stage of the central Parate- upper Quaternary sequences of the Po Plain, across fl ooding surfaces and a sequence bound- thys: Brno, Czech Republic, Masaryk Univer- Italy: Geology, v. 32, p. 989–992, doi:10.1130 sity, p. 107–109. /G20808.1. ary may support the hypothesis of homeostasis Harzhauser, M., Böhme, M., Mandic, O., and Hof- Van Wagoner, J.C., Mitchum, R.M., Campion, K.M., formulated for modern estuaries. mann, C.-C., 2002, The Karpatian (Late Bur- and Rahmanian, V.D., 1990, Siliciclastic se- digalian) of the Korneuburg Basin—A palaeo- quence stratigraphy in well logs, cores, and ACKNOWLEDGMENTS ecological and biostratigraphical synthesis: outcrops: Concepts for high-resolution correla- We thank S. Ćorić, K. Kleemann, A. Kroh, K. Beiträge zur Paläontologie, v. 27, p. 441–456. tion of time and facies: American Association Rauscher, O. Schultz, and W. Sovis for help with Holland, S.M., 2000, The quality of the fossil re- of Petroleum Geologists Methods in Explora- fi eld work and species identifi cations; P. Strauss for cord: A sequence stratigraphic perspective: tion Series 7, 63 p. help with the sequence stratigraphic interpretation; Paleobiology, v. 26, supplement, p. 148–168, Wessely, G., 1998, Geologie des Korneuburger Beck- S. Dominici, M. Stachowitsch, and A. Tomasovych doi:10.1666/0094-8373(2000)26[148:TQOTF ens: Beiträge zur Paläontologie, v. 23, p. 9–23. for discussions; and C.E. Brett, S. Holland, and M. R]2.0.CO;2. Whitfi eld, A.K., and Elliott, M., 2011, Ecosystem and Kowalewski for stimulating reviews. This study was Holland, S.M., and Patzkowsky, M.E., 2004, Ecosys- biotic classifi cation of estuaries and coasts, in supported by project P19013-B17 of the Austrian tem structure and stability: Middle Upper Ordo- Wolanski, E., and McLusky, D.S., eds., Treatise Science Fund (FWF). vician of central Kentucky, USA: Palaios, v. 19, on estuarine and coastal science: Amsterdam, p. 316–331, doi:10.1669/0883-1351(2004)019 Elsevier, p. 99–124. REFERENCES CITED <0316:ESASMU>2.0.CO;2. Wolff, W.J., 1983, Estuarine benthos, in Ketchum, Boyd, R., Dalrymple, R.W., and Zaitlin, B.A., 2006, Holterhoff, P.F., 1996, Crinoid biofacies patterns in B.H., ed., Estuaries and enclosed seas: Eco- Estuarine and incised-valley facies models, in Upper cyclothems, midcontinent, systems of the world 26: Amsterdam, Elsevier, Posamentier, H.W., and Walker, R.G., eds., North America: The role of regional processes p. 151–182. Facies models revisited: SEPM (Society for in biofacies recurrence: Palaeogeography, Pal- Sedimentary Geology) Special Publication 84, aeoclimatology, Palaeoecology, v. 127, p. 47– Manuscript received 24 May 2013 p. 171–235, doi:10.2110/pec.06.84.0171. 81, doi:10.1016/S0031-0182(96)00088-0. Revised manuscript received 5 September 2013 Brett, C.E., Hendy, A.J.W., Bartholomew, A.J., Latal, C., Piller, W.E., and Harzhauser, M., 2006, Manuscript accepted 9 September 2013 Bonelli, J.R., and McLaughlin, P.I., 2007, Re- Small-scaled environmental changes: Indica- sponse of shallow marine biotas to sea-level tions from stable isotopes of gastropods (Early Printed in USA

10 www.gsapubs.org | January 2014 | GEOLOGY GSA DATA REPOSITORY 2014002 M. Zuschin et al.

Table DR1. Abundance and species richness of higher taxa in the studied quantitative samples.

Higher taxon number of families number of species number of fossils Demospongea 1 1 166 Scleractinia 3 4 540 Polychaeta 1 1 19 Cirripedia 1 2 395 Decapoda 1 1 34 Molluscs 62 138 36122 Echinodermata 1 1 3 Chondrichthyes 3 3 34 Osteichthyes 6 8 232 Total 80 159 37545

Table DR2. Number of samples in the four biofacies along with their environmental and stratigraphic affiliation.

Cluster 1 Cluster 2 Cluster 3 Cluster 4 Biofacies Agapilia Granulolabium- Nassarius-Turritella- Nassarius- Agapilia Corbula Paphia-Loripes Total no. of samples 20 31 28 29 Water depth intertidal 19 31 2 8 subtidal 1 0 26 21 Sediment type pelitic 14 17 23 6 sandy 6 14 5 23 Position W 17 16 25 19 E 3 9 3 10 Swell 0 6 0 0 Sequence one 15 25 14 22 two 5 0 14 7 Systems tract TST1 2 23 11 20 HST 13 8 4 2 TST2 5 0 13 7

Figure DR 1. The proximal section Stetten East consists of TST1 annd the lowermost part of the HST. It can be correlated by lithology and faunal content wwith the lower part of section Stetten West (see Fig. 1).

Figure DR2. Two-way cluster analysis of benthic assemblages.

Figure DR3: Ordination plots of non-metric multidimensional scaling (nMDS). A. Distribution of samples in relation to systems tracts. B. Distribution of samples in relation to grain size of sediments. C. Distribution of samples in relattion to biofacies.

Figure DR4: The most abundant taxa in the four biofacies (after correction of data for disarticulation of bivalves).

Cluster analysis

Q-mode cluster analysis resulted in four well-defined clusters, each representing a distinct biofacies (Fig. DR2). Cluster A (20 samples) is characterized by the Agapilia biofaciees (Fig. DR4) and consists almmost exclusively of samples from intertidal settings, more than two thirds are from pelitic sediments and three quarters are from the lower fourth order cycle. Two thirds of the samples here are from the HST, the rest belongs mostly to TST2 (Table DR2). Cluster 2 (31 samples) is characterized by the Granulolabium-Agapilia biofacies (Fig. DR2, Fig DR4). It consists of intertidal samples only, all belong to the lower fourth order cycle (including all samples from the Flysch swell), three quarters are from TST1, the rest from the HST, and almost half of the samples are from sandy sediments (Table DR2). Cluster 3 (28 samples) is characterized by the Nassarius-Turritella-Corbula biofacies (Fig. DR2, Fig. DR4). Virtually all samples here were classified as sublittoral, the great majority is from pelitic sediments and half belongs to the upper fourth order cycle. The majority of samples belong in roughly equal numbers to the two TSTs, only four are from the HST (Table DR2). Cluster 4 (29 samples) is characterized by the Nassarius-Paphia- Loripes biofacies (Fig. DR2, Fig. DR4). The majority of samples here is from sandy, subtidal settings and three quarters belong to the lower fourth order cycle. Samples are mostly from TST1, followed by TST2, only two are from the HST (Table DR2). R-mode cluster analysis revealed four clusters of taxa with similar distributions among samples (Fig. DR2).Cluster I consists of Agapilia and Granulolabium, two species of gastropods that strongly dominate cluster A (Agapilia biofacies) and B (Granulolabium-Agapilia biofacies) and are considered as indicators of intertidal environments. Cluster II consists of the gastropods Nassarius and Turritella, and the bivalves Corbula, Striarca, Pelecyora and Anadara, which are all typical components of cluster C (Nassarius-Turritella-Corbula biofacies) and can be considered as indicators of sublittoral and rather pelitic environments. Cluster III consists of Loripes and Paphia, two bivalve species which typically occur in samples of cluster D (Nassarius-Paphia-Loripes biofacies) and are considered as indicators of sublittoral sandy environments. Cluster IV consists of the remaining species, which are best characterized by their rather patchy distributions. Accordingly, most of these species occur in low mean abundances (<5%) in all biofacies. Exceptions are the coral Porites, which is abundant in cluster 2 and Bittium, which is abundant in cluster 1.

Appendix DR1: Actualistic comparison of abundant taxa

The quantitatively most important taxon Agapilia is an extinct member of the neritid gastropods, a family which today is present in different habitats worldwide but mostly in intertidal rock and mangrove regions (subtropical and tropical) (Scott and Kenny, 1998). The quantitatively very important Granulolabium belongs to the potamidid gastropods, which today also live in the intertidal, in estuaries or sandy mudflats (Healy and Wells, 1998) and are known from temperate and tropical mudflats in the Indo-West Pacific (Ewers, 1963; Wells, 1984). Nassariid gastropods which occur globally from intertidal to shallow subtidal marine habitats (Cernohorsky, 1984) but mostly in estuarine and shallow marine (soft-substrate) environments of temperate and tropical zones (Harasewych, 1998) are also numerous. The two abundant species of this family in our study, Nassarius edlaueri and Cyllenina ternodosa were also recorded from coeval deposits from northern Korneuburg basin and are there interpreted as brackish-marine species, dwelling on intertidal mudflats to shallow sublittoral habitats (Zuschin et al., 2004). Cerithiid gastropods (“sand creepers”) are common in shallow marine water in subtropical and tropical zones globally and in different substrata like sand flats or mangroves (Healy and Wells, 1998), and the Bittium mainly occurs in vegetated marine environments (Bernasconi and Stanley, 1997; Olabarria et al., 1998; Schneider and Mann, 1991, Weber & Zuschin 2013). The brackish and freshwater species Hydrobiidae occur in different habitats like rivers or estuarine mudflats (Ponder and De Keyzer, 1998); they are for example typical components of inner tidal flat assemblages in the northern Adriatic Sea (Weber & Zuschin, 2013) and of tidal flats on the Atlantic coast of France (Poirier et al. 2010). Turritellidae are mostly common on sublittoral muddy-sandy bottoms (Healy and Wells, 1998); they were for example abundantly present in sublittoral muddy sediments of the northern Adriatic Sea (Sawyer & Zuschin 2010). Scaliolid gastropods like Sandbergeria perpusilla live in sandy-mud substrate in the intertidal and littoral (Healy and Wells, 1998) and are known from the subtropical and tropical Indo-West Pacific (Ponder, 1994). In the northern Red Sea, scaliolids are widely distributed, but most abundant in sublittoral muddy sediments (Janssen et al. 2011). The most abundant bivalve in our study is Loripes, a member of the Lucinidae which are known from the intertidal to the shallow subtidal (Reid and Slack-Smith, 1998; Taylor and Glover, 2006), typically harbour chemosymbionts (Berg and Alatalo, 1984; Reid and Brand, 1986; Reid and Slack-Smith, 1998; Johnson & Fernandez 2001) and are often associated with seagrass environments in warm waters (Barnes and Hickman, 1999; Johnson et al., 2002, Zuschin & Oliver 2003, van der Heide et al. 2012). The venerid bivalve Paphia occurs worldwide in different environments but mostly in the intertidal of tropical (Indo-Pacific) to temperate regions (Harte, 1998) and is known from subtidal habitats in the Mediterranean and the Red Sea (e.g. Poppe and Goto 1993, Zuschin and Oliver 2003). Corbulidae are shallow subtidal inhabitants in soft-bottom environments (Hrs-Brenko, 2006; Lamprell et al., 1998) and Corbula gibba lives in the coastal and estuarine subtidal (Holmes and Miller, 2006). Cardiidae (“heart cockles”) live mostly infaunal in shallow habitats (Rufino et al., 2010, Wilson, 1998) and the genus Cerastoderma prefers estuarine conditions in temperate regions (Boyden and Russel, 1972). Living Cerastoderma in the northern Adriatic Sea occur from the outer tidal flat to the shallow sublittoral, but its empty valves are frequently distributed across the tidal flat (Weber & Zuschin 2013). On the Atlantic coast of France, living Cerastoderma is restricted to the tidal flats, but its empty valves are transported into the sublittoral (Poirier et al. 2010). Striarca is a neotiid bivalve, and this family is today common in the intertidal and shallow sub- littoral (Stanley, 1970; Oliver, 1985). Striarca lactea occurs today in the sublittoral northern Adriatic Sea in shallow, sandy sediments (Weber & Zuschin 2013). The endobyssate arcid bivalve Anadara today lives in sublittoral environments, for example in the northern Red Sea (Zuschin & Oliver 2003) and northern Adriatic Sea (Sawyer & Zuschin 2010). Porites is very abundant in Miocene reefs of the Paratethys Sea (e.g., Riegl & Piller 2000, Reutter & Piller 2011) and among the main builders in modern coral reefs (e.g. Cortés et al. 1994, Grossman and Fletcher 2004, Macintyre et al. 1992). Quantitatively less important but environmentally indicative are clionids, balanids, decapods and goniasterids. Clionids are mainly limestone- boring sponges (Goreau and Hartman, 1963; Rützler, 1975) and often dominate shallow-water sponge associations (Rützler, 2002). Balanus amphitrite is an acorn barnacle (Desai and Anil, 2005) which lives in the intertidal (Thiyagarajan, 2010). The recent B.amphitrite is known from warm, tropical and Balanus tinntinabulum from temperate and tropical regions (Wöhrer, 1998). The unspecified decapod-claws could indicate a marine, soft-muddy environment (Müller, 1998). Goniasterid star fishes typically occur in shallow sublittoral habitats (Villier et al., 2004).

References

Barnes, P.A.G., Hickman, C.S., 1999. Lucinid bivalves and marine angiosperms: a search for causal relationships. In: Walker, D.I., Wells, F.E. (Eds.), The Seagrass Flora and Fauna of Rottnest Island, Western Australia. Western Australian Museum, Perth, 215-238. Berg, C.J., Alatalo, P., 1984. Potential of chemosymbiosis in molluscan mariculture. Aquaculture 39, 165-179. Bernasconi, M.P., Stanley, D.J., 1997. Molluscan biofacies, their distributions and current erosion on the Nile delta shelf. Journal of Coastal Research 13, 1201- 1212. Boyden, C.R. & Russel, P.J.C., 1972. The distribution and habitat range of the brackish water cockle (Cardium (Cerastoderma) edule) in the British Isles. Journal of Ecology, 41, 719-734. Cernohorsky, O., 1984. Systematics of the family Nassariidae. Bulletin of the Auckland Institute and Museum 14, 356. Cortés, J., Macintyre, I.G., Glynn, P.W., 1994. growth history of an eastern Pacific fringing reef, Punta Isotes, Costa Rica. Coral Reefs 13, 65-73. Desai, D.V., Anil, A.C., 2005. Recruitment of the barnacle Balanus amphitrite in a tropical estuary: implications of environmental perturbation, reproduction and larval ecology. Journal of the Marine Biological Association of the United Kingdom 85, 909-920. Ewers, W.H., 1963. Some observations on the life cycle of Velacumantus australis (Quoy & Gaimard) (Gastropoda: Potamididae). Journal of the Malacological Society of Australia 7, 62-71. Goreau, T.F., Hartman, W.D., 1963. Boring sponges as controlling factors in the formation and maintenance of coral reefs. In: Sognnaes, R.F. (Ed.), Mechanisms of Hard Tissue Destruction. Publications of the American Association for the Advancement of Science 75, 25-54. Grossman, E.E., Fletcher, C.H., 2004. Holocene reef development where wave energy reduces accommodation space, Kailua Bay, windward Oahu, Hawaii, U.S.A. Journal of Sedimentary Research 74, 49-63. Harasewych, M.G., 1998. Family Nassariidae. In: Beesley, P.L., Ross, G.J.B., Wells, A (Eds.), - The Southern Synthesis: Fauna of Australia, Part B, 5, 829- 831. Harte, M.E., 1998. Superfamily Veneroidea. In: Beesley, P.L., Ross, G.J.B., Wells, A. (Eds.), Mollusca - The Southern Synthesis: Fauna of Australia, Part A, 5, 355- 362. Healy, J.M., Wells, F.E., 1998. Superfamily Cerithioidea. In: Beesley, P.L., Ross, G.J.B., Wells, A. (Eds.), Mollusca - The Southern Synthesis: Fauna of Australia, Part B, 5, 707-733. Holmes, S.P., Miller, N., 2006. Aspects of the ecology and population genetics of the bivalve Corbula gibba. Marine Ecology Progress Series 315, 129-140. Hrs-Brenko, M., 2006. The basket shell, Corbula gibba Olivi, 1792 (Bivalve Mollusks) as a species resistant to environmental disturbances: A review. Acta Adriatica 47(1), 49-64. Janssen, R., Zuschin, M. Baal, Ch., 2011, Gastropods and their habitats from the northern Red Sea (Egypt: Safaga) Part 2: Caenogastropoda: Sorbeoconcha and Littorinimorpha. Annalen des Naturhistorischen Museums Wien 113A, 373- 509. Johnson, M.A., Fernandez, C., Pergent, G., 2002. The ecological importance of an invertebrate chemoautotrophic symbiosis to phanerogam seagrass beds. Bulletin of Marine Science 71, 1343-1351. Lamprell, K., Healy, J.M., Dyne, G.R., 1998. Superfamily Myoidea. In: Beesley, P.L., Ross, G.J.B., Wells, A. (Eds.), Mollusca - The Southern Synthesis: Fauna of Australia, Part A, 5, 363-366. Macintyre, I.G., Glynn, P.W., Cortés, J., 1992. Holocene Reef History in the Eastern Pacific: Mainland Costa Rica, Caño Island, Cocos Island, and Galápagos Islands. Proceedings of the 7th International Coral Reef Symposium 2, 1174- 1184. Müller, P., 1998. Decapode Crustacea aus dem Karpat des Korneuburger Beckens (Unter-Miozän, Niederösterreich). Beiträge zur Paläontologie 23, 273-281. Olabarria, C., Urgorri, V., Troncoso, J.S., 1998. An analysis of the community structure of subtidal and intertidal benthic mollusks of the Inlet of Bano (Ria de Ferrol) (northwestern Spain). American Malacological Bulletin 14, 103-120. Oliver, P.G., 1985. A comparative study of two species of Striarciinae from Hong Kong with comments on specified and generic systematics. In: Morton, B., Dudgeon, D. (Eds.), Proceedings of the Second International Workshop on the Malacofauna of Hong Kong and Southern China, Hong Kong University Press, 283-310. Poirier, C. Sauriau, P-G., Chaumillon, E. Bertin, X. 2010, Influnce of hydro- sedimentary factors on mollusk death assemblages in a temperate mixed tide- and wave- dominated coastal environment: Implications for the fossil record. Continental Shelf Research 30, 1876-1890. Ponder, W.F., 1994. The anatomy and relationships of Finella and Scaliola (Caenogastropoda: Cerithiodea: Scaliolidae). In: Morton, B. (Ed.), The Malacofauna of Hong Kong and Southern China. Proceedings of the Third International Workshop on the Malacofauna of Hong Kong and Southern China, Hong Kong University Press, 215-241. Ponder, W.F., De Keyzer, R.G., 1998. Superfamily Rissoidea. In: Beesley, P.L., Ross, G.J.B., Wells, A. (Eds.), Mollusca - The Southern Synthesis: Fauna of Australia, Part B, 5, 745-766. Poppe, G.T., Goto, Y., 1993. European seashells, Volume 2. Verlag Christa Hemmen, Wiesbaden. Reid, R.G.B., Brand, D.G., 1986. Sulfide-oxidising symbiosis in lucinaceans: implications for bivalve evolution. Veliger 29, 3-24. Reid, R.G.B., Slack-Smith, S.M., 1998. Superfamily Lucinoidea. In: Beesley, P.L., Ross, G.J.B., Wells, A. (Eds.), Mollusca - The Southern Synthesis: Fauna of Australia, Part A, 5, 309-315. Riegl, B. & Piller, W.E. 2000, Biostromal Coral Facies—A Miocene Example from the Leitha Limestone (Austria) and its Actualistic Interpretation. Palaios 15, 399-413 Reutter, M. & Piller, W.E. 2011, Volcaniclastic events in coral reef and seagrass environments: evidence for disturbance and recovery (Middle Miocene, Styrian Basin, Austria). Coral Reefs 30, 889-899 Rützler, K., 1975, The role of burrowing sponges in bioersion. Oecologia, v.19, 203- 216. Rützler, K., 2002. Impact of crustose clionid sponges on Caribbean reef corals. Acta Geologica Hispanica 37, 61-72. Rufino, M.M., Gaspar, M.B., Pereira, A.M., Maynou, F., Monteiro, C.C., 2010. Ecology of megabenthic bivalve communities from sandy beaches on the south coast of Portugal. Scientia Marina 74(1), 163-178. Sawyer, J.A, Zuschin, M. 2010, Intensities of drilling predation from molluscan assemblages along a transect through the northern Gulf of Trieste ( Adriatic Sea ). Palaeogeography, Palaeoclimatology, Palaeoecology 285, 152-173. Schneider, F.I., Mann, K.H., 1991. Species specific relationship of invertebrates to vegetation in a seagrass bed: II. Experiments on the importance of macrophyte shape, epiphyte cover and predation. Journal of Experimental Marine Biology and Ecology 145, 119-139. Scott, B.J., Kenny, R., 1998. Superfamily Neritoidea. In: Beesley, P.L., Ross, G.J.B., Wells, A. (Eds.), Mollusca - The Southern Synthesis: Fauna of Australia, Part B, 5, 694-702. Stanley, S.M., 1970. Relation of the shell form to life habits of the (Mollusca). Memoirs, Geological Society of America 125, 1-296. Taylor, J.D., Glover, E.A., 2006. Lucinidae (Bivalvia) - the most diverse group of chemosymbiotic molluscs. Zoological Journal of the Linnean Society 148, 421- 438. Thiyagarajan, V., 2010. A review on the role of chemical cues in habitat selection by barnacles: New insights from larval proteomics. Journal of Experimental Marine Biology and Ecology 392, 22-36. Van der Heide, T., Govers, L.L., de Fouw, J., Olff, H., van der Geest, M. van Katwijk, M.M., Piersma, T., van de Koppel, J., Silliman, B.R., Smolders, A.J.P., van Gils, J.A. 2012, A three-stage symbiosis forms the foundation of seagrass ecosystems. Science 336, 1432-1434 Villier, L., Kutscher, M., Mah, Ch.L., 2004. Systematics and palaeoecology of middle Toarcian Asteroidea (Echinodermata) from the “Seuil du Poitou”, Western France. Geobios 37, 807-825. Weber, K, Zuschin, M. 2013, Delta-associated molluscan life and death assemblages in the northern Adriatic Sea: Implications for palaeoecology, regional diversity and conservation. Palaeogeography, Palaeoclimatology, Palaeoecology 370, 77-91. Wells, F.E., 1984. Population characteristics of the periwinkle, Nodilittorina unifasciata, on a vertical rock clif in Western Australia. Nautilus 98, 102-107. Wilson, B., 1998. Superfamily Cardioidea. In: Beesley, P.L., Ross, G.J.B., Wells, A. (Eds.), Mollusca - The Southern Synthesis: Fauna of Australia, Part A, 5, 328- 332. Wöhrer, S., 1998. Balanidae (Crustacea, Cirripedia) aus dem Karpat des Korneuburger Beckens. Beiträge zur Paläontologie 23, 283-293. Zuschin, M., Oliver, G., 2003. Bivalves and bivalve habitats in the northern Red Sea - The Northern Bay of Safaga (Red Sea, Egypt): An actuopalaeontological approach, VI. Bivalvia, Naturhistorisches Museum, Wien. Zuschin, M., Harzhauser, M., Mandic, O., 2004. Palaeoecology and taphonomy of a single parautochthonous Paratethyan tidal flat deposit (Karpatian,Lower Miocene - Kleinebersdorf, Lower Austria). Courier Forschungsinstitut Senckenberg 246, 153-168.