Document [original]

Influence of abiotic and biotic factors on benthic marine

community composition, structure and stability: a

multidisciplinary approach to molluscan assemblages from the

Miocene of northern Germany

by DARIA CAROBENE

1,2,3,

*, ROBERT BUSSERT

,ULRICHSTRUCK

CARL J. REDDIN

and MARTIN ABERHAN

LWL-Museum of Natural History, Westphalian State Museum with Planetarium, Sentruper Straße 285, 48161 M€

unster, Germany; daria.carobene@lwl.org

Museum f€

ur Natur und Umwelt L€

ubeck, M€

uhlendamm 1–3, 23552 L€

ubeck, Germany

Museum f€ur Naturkunde, Leibniz-Institut f€ur Evolutions- und Biodiversit€atsforschung, Invalidenstraße 43, 10115 Berlin, Germany

Institut f€

ur Angewandte Geowissenschaften, Technische Universit€

at Berlin, Ernst-Reuter-Platz 1, 10587 Berlin, Germany; [email protected]

*Corresponding author

Typescript received 28 September 2022; accepted in revised form 22 March 2023

Abstract: The Miocene mica-clay deposits of Groß Pam-

pau (northern Germany) are well known for their diverse

assemblages of marine mammals. Despite numerous system-

atic and biostratigraphic studies, an in-depth palaeoecologi-

cal analysis of its molluscan assemblages and a

comprehensive palaeoenvironmental reconstruction are lack-

ing. Here, we integrate new faunal, sedimentological and

geochemical data to reconstruct the marine palaeoecosystem

of the Upper Miocene sedimentary succession of Groß Pam-

pau, and to identify the drivers controlling the composition,

ecological structure and temporal dynamics of its macro-

benthic molluscan assemblages. Fossil evidence, coupled with

analyses of clay mineral composition, grain size distribution

and geochemical data (total organic carbon, total nitrogen,

C, d

O, d

N of sediment and shells), suggests a warm–

temperate, mesotrophic, low-energy, offshore marine setting

mostly below storm wave base and a pronounced surface-to-

bottom water temperature gradient. Low variability in sedi-

mentological and geochemical signals indicates generally

stable physicochemical conditions, whereas the occurrence of

the opportunistic species Varicorbula gibba suggests occa-

sionally unfavourable bottom conditions, possibly related to

transient hypoxia. Canonical correspondence analysis indi-

cates that the distribution of molluscan assemblages corre-

lates with total organic carbon and nitrogen content,

suggesting organic matter availability at the sea floor as a

controlling factor. A pattern of repetitive punctuated stasis

of molluscan assemblages is defined by the temporal persis-

tence in taxonomic and ecological composition, occasionally

interrupted by shifts to a different faunal configuration. We

suggest that both stable environmental conditions and biotic

interactions (i.e. the top-down control exerted by carnivo-

rous gastropods and environmental modification by ubiqui-

tous burrowing deposit feeders) probably contributed to the

observed temporal stability.

Key words: Late Miocene, North Sea Basin, palaeoenviron-

ment, molluscan fauna, stable isotopes.

OFTEN referred to as ‘the whales’ graveyard’, the locality

of Groß Pampau in northern Germany is famous for its

fossil whale remains. The fossils are derived from fully

marine mica-clay deposits, which developed in the south-

east of the North Sea Basin during the Middle–Late

Miocene, and are found today at Groß Pampau, in the

commercial gravel and clay pit of the company Kieswerke

Ohle & Lau GmbH (Hinsch 1990). Over the past

40 years, excavations yielded highly diverse marine fossil

assemblages, which include marine mammals (mainly

mysticetes, odontocetes and pinnipeds), remains of chelo-

niid and dermochelyid sea turtles, teeth and skeletal

elements of elasmobranchs, teleost otoliths, and more

than 140 species of mollusc (Moths 1989,1990,1992,

1994,1995,1998,2003;H

€

opfner 1991,2014; Behrmann

1995; Lierl 1995; Hampe 1999,2006; Kaz

ar & Hampe

2014; Kriwet et al.2015; Monta~

nez-Rivera & Hampe

2020).

Despite numerous studies on the systematics and bio-

stratigraphy of the Groß Pampau succession, an in-depth

reconstruction of the Late Miocene marine palaeoenvir-

onment and of the importance of abiotic and biotic

parameters that primarily governed the composition and

dynamics of its species-rich molluscan assemblages is

Papers in Palaeontology published by John Wiley & Sons Ltd on behalf of The Palaeontological Association.

doi: 10.1002/spp2.1496 1

This is an open access article under the terms of the Creative Commons Attribution License,

which permits use, distribution and reproduction in any medium, provided the original work is properly cited.

[Papers in Palaeontology, 2023, e1496]

lacking. To fill this gap, we used a multidisciplinary

approach, comprising palaeoecological, sedimentological

and geochemical analyses. Based on new quantitative bed-

by-bed sampling, we analysed the taxonomic composi-

tion, diversity and ecological structure of macrobenthic

assemblages and their dynamic fluctuations through time.

For a regional perspective, we compared the Groß Pam-

pau assemblages with contemporaneous benthic mollus-

can assemblages in the North Sea Basin, in particular

those from the Late Miocene of Denmark (Rasmus-

sen 1966). Coupled with new sedimentological analyses of

grain size distributions and clay mineral composition,

and utilizing geochemical proxy data (total organic car-

bon, total nitrogen, d

C, d

O, and d

N of bulk sedi-

ment and selected shells), we provide a detailed view of

the Late Miocene palaeoenvironmental conditions of the

Groß Pampau area. Specifically, we address the following

questions: (1) which were the prevailing physicochemical

conditions, and how stable or dynamic were these over

time; (2) how are benthic molluscan assemblages charac-

terized in terms of species diversity, taxonomic composi-

tion, and life-habit and trophic structure, and how stable

or variable were they; (3) are there correlative, and thus

potentially causative, links between abiotic ecosystem

components and the characteristics of molluscan assem-

blages; and (4) how important were biotic parameters,

such as predation pressure, for molluscan assemblage

composition and structure and their dynamics.

GEOLOGICAL SETTING

The study area (53°31047.800 N, 10°33055.400 E) is

located in the commercial gravel and clay pit of the com-

pany Kieswerke Ohle & Lau GmbH in Groß Pampau,

northern Germany (Fig. 1). The sedimentary succession

of the Pampau area developed south-east of the North

German Cenozoic troughs in the North Sea Basin and

geographically represents the largest area of pre-

Quaternary outcrops in the state of Schleswig-Holstein

(Hinsch 1990).

The development of the Miocene sedimentary succes-

sion of the North Sea Basin was influenced by tectonic

events and marked climatic changes (Jordt et al.1995;

Clausen et al.1999; Rasmussen 2004a,2004b; Larsson

et al.2011). At the end of the Early Miocene and begin-

ning of the Middle Miocene, the global warming known

as the Miocene Climate Optimum coincided with a dis-

tinct sea level rise and deposition of silts and marine

sands of the Arnum Formation (Hinsch 1986a;

Rasmussen 2004a,2004b; Rasmussen et al.2010). Mean-

while, in the eastern part of the North Sea Basin, tecto-

nism and uplift of the hinterland led to a major increase

of sediment supply and consequent progradation of the

shoreline with the development of the fluvial Upper

Brown Coal Sand Formation (Gripp 1964; Hinsch 1986a;

Rasmussen 2004a,2004b; Gibbard & Lewin 2016). Despite

the global climatic cooling of the Middle Miocene Cli-

mate Transition (13.9 Ma; Westerhold et al.2005), depo-

sition of fine-grained marine sediments (Hodde and

Gram formations) dominated the late Middle Miocene to

early Late Miocene as a result of increased subsidence of

the eastern North Sea Basin (Hinsch 1986a;

Rasmussen 2004a,2004b). Progradation of the coastline

continued during the Late Miocene to Early Pliocene,

leading to the deposition of shoreface sediments (Morsum

Member) and fluvial deposits (Oldesloe Formation)

(Hinsch 1986a; Rasmussen 2004a,2004b). In the Groß

Pampau pit, below c.1–3-m-thick Pleistocene deposits, c.

17-m-thick mica-rich clay and silt layers of the so-called

‘Oberer Glimmerton’ (Upper Mica-Clay Formation) are

exposed (Spiegler & G€

urs 1996). Two cored sections

(Pampau I and II), drilled in the pit by the Geological

Survey of Schleswig-Holstein, showed that the fully

marine deposits developed during the late Langhian

(Middle Miocene) to early Tortonian (Late Miocene)

(Hinsch 1990; Spiegler & G€

urs 1996). According to litho-

and biostratigraphical comparisons with the adjacent

southern Denmark successions, the lower and upper

FIG. 1. Palaeogeographical reconstruction of west-central

Europe during the Middle–Late Miocene showing the geographi-

cal position of the localities of Groß Pampau and Gram (modi-

fied from Kriwet et al.2015).

2PAPERS IN PALAEONTOLOGY

deposits of the ‘Oberer Glimmerton’ of Schleswig-

Holstein are usually referred to respectively as the Hodde

and the Gram formations. As a result of transgressive

events, the boundary between the Hodde and Gram for-

mations is diachronous and corresponds approximately to

a glauconitic clay unit in the upper part of the cored sec-

tion Pampau I (King et al.2016).

The cores were studied biostratigraphically using ben-

thic and planktonic molluscs (Hinsch 1990; Spiegler &

G€

urs 1996;G

€

urs 2002;G

€

urs & Janssen 2002) and bolbo-

forms (Spiegler & G€

urs 1996;G

€

urs & Spiegler 1999). The

benthic molluscan assemblages define the regional sub-

stages of the upper Reinbekian to upper Langenfeldian

(Hinsch 1990; Spiegler & G€

urs 1996). The Bolboforma

biostratigraphy enabled an accurate calibration of the

regional stages, and correlation with the nannoplankton

(Spiegler & G€

urs 1996) and the planktonic gastropod bio-

zonations (G€

urs & Janssen 2002). Additional biostratigra-

phical studies on the Groß Pampau boreholes and

sedimentary succession included calcareous nannoplank-

ton (Martini 2001), and dinoflagellate cysts and forami-

nifera (Daniels et al.1990). However, due to the absence

of key species (Martini 2001) and lack of stratigraphical

order in sampling (Daniels et al.1990), correlation with

the studied section remains unresolved.

The succession studied herein corresponds to the lower

part of the Gram Formation and biostratigraphically cor-

relates with the upper 5–6 m of the borehole Pampau I,

as presented in Hinsch (1990). The presence of Astarte

vetula and Astarte gleuei in the lower and upper part of

the section, respectively, correlates with the upper Lan-

genfeldian substage (sensu Hinsch 1986b,1990) and the

Bolboforma fragori–subfragoris and B. capsula–laevis zones.

Occurrence of the planktonic gastropods Limacina valva-

tina,Limacina gramensis and Limacina ingridae support

the biostratigraphic inferences based on benthic molluscs.

In particular, the presence of L. ingridae and the concom-

itant absence of L. gramensis in the upper 2 m restrains

this part of the section to the B. capsula–laevis Zone

(G€

urs & Janssen 2002). In conclusion, based on the ben-

thic and planktonic molluscs and correlation with the

Bolboforma biozonation, we infer an approximately early

Tortonian age for the studied succession.

MATERIAL AND METHOD

Field work

The 6-m-thick sedimentary succession is located in the

north-eastern part of the pit on a south-facing wall

(Fig. 2). To investigate the taxonomic and ecological

composition of the benthic molluscan fauna, 30 bulk

samples of c. 20 kg each were collected from uniform

layers of sandy silt at intervals of 20 cm (Fig. 2). Addi-

tionally, 30 sediment samples of c. 500 g each were taken

for sedimentological and stable isotope analyses from the

same levels as the faunal samples.

Sedimentological analysis

Grain size analysis was carried out with a combined sieve

and areometer analysis. Samples were washed through a

sieve with a 63 lm mesh size. The sand fraction was dry

sieved in quarter-phi steps. The grain size distribution of

the silt and clay fractions was determined by areometer

analysis on sub-samples weighing c.30–50 g. The results of

the combined sieve and areometer analysis were evaluated

using the programs GGU-SIEVE (https://www.ggu-

software.com/geotechnik-software/laborversuche/ggu-sieve-

kornverteilung) and GRADISTAT (Blott & Pye 2001).

To determine the clay mineralogy of the samples, the

clay fractions were separated using a centrifuge and

smeared onto slides. Each sample was measured three

times in a Bruker D2 Phaser x-ray diffractometer with

Cu-Karadiation from 3 to 300 2h, first after air drying,

then after ethylene glycol treatment and finally after heat-

ing at 550°C for 1 h. Clay minerals were identified by

comparing the three x-ray diffractometer runs. Quantifi-

cation of the clay mineralogy was performed on the ethyl-

ene glycol runs, with the areas of mineral-specific peaks

measured with Rigaku’s SmartLab software and converted

to relative weight fractions using the weighting factors of

Biscaye (1965). We also calculated the clay mineral ratio

illite +kaolinite/smectite +chlorite (IK/SC). Variation in

this ratio can indicate shifting contributions of clay min-

erals from different source areas, climatic variations such

as those induced by orbital forcing (e.g. Milankovitch

cycles), or a combination of both.

Faunal samples

All samples were dried, soaked in water and washed over

a 0.5 mm mesh sieve. Residues were treated with 10%

solution and wet-sieved through a 0.5 mm mesh

screen. Taxa were identified to the lowest possible taxo-

nomic rank (species or genus). The total dataset consists

of 6810 individuals representing 112 benthic taxa of gas-

tropods, bivalves, scaphopods and one brachiopod (Caro-

bene et al.2023, datasheet 1). The benthic fauna

occasionally included remains of echinoids and cupulidrid

bryozoans. Owing to their fragmentary condition, it was

unclear how many complete specimens they represented

and thus were not considered further in the quantitative

analyses. A total of 3090 occurrences of three planktonic

gastropod species were used for biostratigraphical

CAROBENE ET AL.: PALAEOENVIRONMENT OF GROß PAMPAU 3

placement. We inferred the number of individuals of

bivalves and brachiopods by adding the most numerous

valves (left–right or dorsal–ventral respectively) to the

number of articulated specimens. For gastropods, com-

plete specimens and individual apices were counted as

single individuals. For scaphopods, the number of indi-

viduals was estimated by counting the number of discrete

posterior apertures. To assess whether the faunal assem-

blages are likely to be (par-)autochthonous relics of for-

mer living communities and shells buried in their original

habitat, we also calculated the percentage of articulated

shells of the six most abundant bivalve species in samples

in which a species reached at least 20 individuals

(Table 1). Specimens were deposited at the Museum f€

Natur und Umwelt L€

ubeck under the sample inventory

numbers MNUL.Pal.100.725–754.

Taxonomy

Taxonomic identifications were based on previous studies

(Moths 1989,1990,1992,1994,1995; Spiegler &

G€

urs 1996) and relevant literature on stratigraphically

coeval and/or geographically close deposits (Ravn 1907;

Kautsky 1925; Gilbert 1945; Hinsch 1952; Rasmussen 1956,

1968; Sorgenfrei 1958; Anderson 1964; Janssen 1984;

FIG. 2. Field photograph and stratigraphy of the Upper Miocene clay pit of Groß Pampau. Photograph provides view of the north-

eastern part of the Ohle & Lau clay pit with sampled section. Note that sampling proceeded from top to bottom, which is reflected in

the numbering of the samples. White arrowheads at top left and bottom right indicate the position of quantitative palaeoecological

sample S1 and S30, respectively. Stratigraphic log of the sampled section shows position of faunal and sediment samples (black arrow-

heads). Abbreviations: c, coarse; f, fine; m, medium; vc, very coarse; vf, very fine. Scale bar represents 1 m.

4PAPERS IN PALAEONTOLOGY

Wienrich 1999,2001,2007; Schnetler 2005; Janssen &

Wienrich 2007;Mothset al.2010;Steinet al.2016). We

used open nomenclature qualifiers in the case of poor

preservation, insufficient number of individuals to evaluate

morphological variability or uncertainty of taxonomic sta-

tus. The taxonomic position of several groups of the Groß

Pampau molluscan fauna will be clarified in a forthcoming

revision (R. Janssen & G. Stein, pers. comm. 2020).

Ecological categorization

We categorized the benthic taxa according to three funda-

mental ecological parameters: mobility level, tiering posi-

tion relative to the sediment–water interface, and feeding

mechanism (Bambach et al.2007; Table 2). We also

assign their specific mode of life (MOL): taxon-specific

unique combinations of these three ecological parameters

that describe the realized ecospace of a community (Bam-

bach et al.2007; Carobene et al.2023, datasheet 2). The

ecological classification is based on the Neogene Marine

Biota of Tropical America (NMITA) database (https://

nmita.rsmas.miami.edu/), the Paleobiology Database

(https://paleobiodb.org/) and the available literature

(Table 2). Out of 112 taxa, only one species (the gastro-

pod Diaphana moerchi represented by a single specimen)

could not be categorized ecologically and was excluded

from the MOL-based analyses. Before performing cluster

analyses, MOLs represented by only one or two individ-

uals were removed from the database because they have

no ecological relevance (non-motile semi-infaunal byssate

suspension feeders: Modiolula cf. phaseolina, one speci-

men; non-motile epifaunal pedically attached suspension

feeders: Cryptopora nysti, two specimens).

Statistical analysis

To investigate taxonomical and functional biodiversity

through time, we calculated richness and Simpson’s index

of diversity for each faunal assemblage. To standardize for

differences in sample size, species richness was calculated

as rarefied richness, with samples rarefied to 91 individ-

uals (i.e. the size of the smallest faunal sample) and using

the rarefy function in the vegan package in R (Oksanen

et al.2019). Simpson’s index considers both richness and

evenness and is defined as 1D, where D =Σp

and p is

the proportional abundance of species i. Simpson’s index

was computed with the diversity function in vegan.

To detect recurring faunal and MOL associations, we

performed two-way cluster analyses. Q-mode hierarchical

clustering (hclust function in the stats package) identifies

groups of faunal assemblages with similar composition

and abundance of species or MOLs, here termed faunal

associations and MOL associations, respectively. R-mode

clustering groups the species or MOLs that tend to co-

occur. Clustering was performed on Bray–Curtis distance

matrices (vegdist function in vegan) using Ward’s cluster-

ing criterion. We applied the similarity profile test (SIM-

PROF) to determine the number of statistically distinct

clusters, calculated with the R package clustsig (Clarke

et al.2008) and used the default significance level of

p<0.05 unless stated otherwise. Illustrations of the two-

way cluster analyses applied the pheatmap() function of

the pheatmap R package. Data were also ordinated using

non-metric multidimensional scaling (NMDS) to verify

the groupings of species and MOLs identified with cluster

analyses. Ordinations were carried out on Bray–Curtis

distance matrices, using the metaMDS function from

vegan, with two dimensions and 100 attempts. The fit of

the NMDS ordinations was estimated using the stress

value: the higher the stress values, the less reliable the

NMDS representation, with values lower than 0.2 consid-

ered acceptable.

To explore the relationship between species occurrence

and the environmental variables measured in this study,

we conducted a canonical correspondence analysis (CCA)

with the function cca (vegan package). The isotopic com-

position of benthic gastropod shells was excluded from

the correspondence analysis because the values were not

normally distributed. The outputs were displayed in a

species-conditional triplot. Sample 5 was excluded from

the final dataset given that no total organic content mea-

surement was available. To minimize the influence of rare

species, the dataset included only species with a total rela-

tive abundance >0.5%. The significance of CCA dimen-

sions and environmental variables was tested using

analysis of variance (ANOVA) with 999 permutations,

applying the function anova.cca (vegan package, Legendre

et al.2011).

We used Spearman’s rank to measure the correlation

between the species accounting for most of the variability

in faunal associations and the environmental variables.

Given that time series can be often spuriously correlated

TABLE 1. Percentage of articulated specimens of the most

abundant Late Miocene bivalve species of Groß Pampau, north-

ern Germany.

Taxon Samples Specimens Articulated specimens

(%)

Yoldiella cf.

pygmaea

25 892 88.79

Yoldiella spiegleri 27 1108 63.63

Astarte gleuei 2 44 9.09

Astarte vetula 4 130 3.08

Alveinus nitidus 8 293 77.82

Varicorbula gibba 7 295 32.20

CAROBENE ET AL.: PALAEOENVIRONMENT OF GROß PAMPAU 5

TABLE 2. Benthic marine macroinvertebrates from the Late Miocene of Groß Pampau, northern Germany and their ecological

classification

†

Class Taxon Mobility Feeding Tiering Remarks and Source

BNucula georgiana Reg-mov Sub-dep Shal-inf Based on Nucula [1;30]

BYoldia cf. glaberrima Reg-mov Sub-dep Shal-inf Based on Yoldia [1;2;43]

BYoldiella cf. pygmaea Reg-mov Sub-dep Shal-inf Based on Yoldiella [1;2;43]

BYoldiella spiegleri Reg-mov Sub-dep Shal-inf Based on Yoldiella [1;2;43]

BModiolula cf. phaseolina Non-mot(bys) Susp Semi-inf Based on Modiolinae [1;10]

BAnadara cf. diluvii Fac-mob(un) Susp Semi-inf Based on Anadara [1;47]

BBathyarca cf. pectunculoides Fac-mob(at) Susp Epi Based on Bathyarca [29]

BLimopsis aurita Fac-mob(at) Susp Semi-inf Based on Limopsis [32]

BKorobkovia sp. Fac-mob(un) Susp Epi Based on Palliolinae [1;4]

BPallolium sp. Fac-mob(un) Susp Epi Based on Palliolum [1]

BPecten sp. Fac-mob(un) Susp Epi Based on Pecten [1;4]

BPseudamussium clavatum Fac-mob(un) Susp Epi Based on Pseudamussium [1;4]

BLimatula sp. Fac-mob(un) Susp Epi Based on Limatula [1;13;47]

BAstarte gleuei Fac-mob(un) Susp Shal-inf Based on Astarte [40;43]

BAstarte vetula Fac-mob(un) Susp Shal-inf Based on Astarte [40;43]

BGoodallia waeli Fac-mob(un) Susp Shal-inf Based on Astarte [40;43]

BCyclocardia sp. Fac-mob(un) Susp Shal-inf Based on Cyclocardia [1]

BAxinulus sp. Fac-mob(un) Chem Deep-inf Based on other Thyasiridae [1;11;22]

BParvicardium cf. scabrum Fac-mob(un) Susp Shal-inf Based on Parvicardium [43]

BAbra cf. antwerpiensis Fac-mob(un) Surf-dep Shal-inf Based on Abra [1;26;50]

BAbra cf. sorgenfreii Fac-mob(un) Surf-dep Shal-inf Based on Abra [1;26;50]

BGlossus olearii Fac-mob(un) Susp Shal-inf Based on Glossus [34]

BAlveinus nitidus Fac-mob(un) Susp Shal-inf Based on Alvenius [1;5]

BSpisula sp. Fac-mob(un) Surf-dep Shal-inf Based on Mactridae [1]

BVaricorbula gibba Fac-mob(at) Susp Shal-inf Based on Corbula [1;49]

BCuspidaria sp. Reg-mov Pred Shal-inf Based on Cuspidaria [1;39]

GTurritellinella tricarinata Reg-mov Susp Semi-inf Based on Turritellidae [1;6;48]

GHemiacirsa lanceolata Reg-mov Par Epi Based on Acirsa [1;17;23;27]

GTurriscala cf. borealis Reg-mov Par Epi Based on other Epitoniidae [1;17;23;27]

GEuspira helicina auct. Reg-mov Pred Shal-inf Based on Euspira [1;7;19]

GNorephora fritschi Reg-mov Par Epi Based on Triphoridae [1]

GAlvania sp. Reg-mov Herb Epi Based on Alvania [1]

GCingula inusitata Reg-mov Herb Epi Based on Cingula [1]

GObtusella gottscheana Reg-mov Herb Epi Based on Obtusella [1]

GSolariorbis pulchralis Reg-mov Herb Epi Based on Solariorbis [1]

GCirculus cf. hennei Reg-mov Herb Epi Based on Circulus [1;41]

GCirculus sp. Reg-mov Herb Epi Based on Circulus [1;41]

GAclis minor Reg-mov Par Epi Based on Acirsa [1;38]

GEulima glabra Reg-mov Par Epi Based on Eulima [1;38]

GMelanella sp. Reg-mov Par Epi Based on other Eulimidae [1;38]

GAporrhais sp. Reg-mov Surf-dep Shal-inf Based on Aporrhais [24;35]

GGaleodea bicatenata marqueti Reg-mov Pred Shal-inf Based on Galeodea [2;3;20;47]

GSemicassis cf. laevigata Reg-mov Pred Shal-inf Based on Semicassis [2;3;20;47]

GXenophora sp. Reg-mov Herb Epi Based on Xenophoridae [1;36]

GCalcarata mioparva Reg-mov Par Epi Based on Cancellariidae [1;33]

GSveltia cf. lyrata Reg-mov Par Epi Based on Cancellariidae [1;33]

GPseudobabylonella cf. fusiformis Reg-mov Par Epi Based on Cancellariidae [1;33]

GPseudobabylonella sp. 1 Reg-mov Par Epi Based on Cancellariidae [1;33]

GPseudobabylonella sp. 2 Reg-mov Par Epi Based on Cancellariidae [1;33]

GPseudobabylonella pampauensis Reg-mov Par Epi Based on Cancellariidae [1;33]

GColus gregarius Reg-mov Pred Epi Based on Colus [1;25;46]

GLiomesus sp. Reg-mov Pred Epi Based on other Buccinidae [1;25;46]

(continued)

6PAPERS IN PALAEONTOLOGY

TABLE 2. (Continued)

Class Taxon Mobility Feeding Tiering Remarks and Source

GMacrurella nassoides Reg-mov Pred Epi Based on Mitrella [1]

GAquilofusus luneburgensis Reg-mov Pred Epi Based on other Fasciolariidae [1;45]

GLatirulus rothi Reg-mov Pred Epi Based on Latirulus [1;45]

GTritia mothsi Reg-mov Sca Epi Based on Nassaridae [1;16]

GTritia sp. 1 Reg-mov Sca Epi Based on Nassaridae [1;16]

GTritia sp. 2 Reg-mov Sca Epi Based on Nassaridae [1;16]

GTritia sp. 3 Reg-mov Sca Epi Based on Nassaridae [1;16]

GTritia cf. spectabilis Reg-mov Sca Epi Based on Nassaridae [1;16]

GTritia syltensis Reg-mov Sca Epi Based on Nassaridae [1;16]

GTritia wienrichi Reg-mov Sca Epi Based on Nassaridae [1;16]

GMurex spinicosta Reg-mov Pred Epi Based on Murex [1;45]

GLyrotyphis cf. sejunctus Reg-mov Pred Epi Based on other Muricidae [1;45]

GPagodula semperi Reg-mov Pred Epi Based on other Muricidae [1;45]

GAmalda cf. obsoleta Reg-mov Pred Epi Based on Olivoidea [44;45]

GConilithes poulsenii Reg-mov Pred Epi Based on Conus [1;45]

GAphanitoma sp. Reg-mov Pred Epi Based on Borsoniidae [1;37;45]

GBathytoma jugleri Reg-mov Pred Epi Based on Bathytoma [1;37;45]

GMicrodrillia serratula Reg-mov Pred Epi Based on Borsoniidae [1;37;45]

GPleurotomoides biconicus Reg-mov Pred Epi Based on Conoidea [1;45]

GPleurotomoides mariae Reg-mov Pred Epi Based on Conoidea [1;45]

GNitidiclavus maitrejus Reg-mov Pred Epi Based on Conoidea [1;45]

GSpirotropis gramensis Reg-mov Pred Epi Based on Conoidea [1;45]

GHaedropleura miocaenica Reg-mov Pred Epi Based on Conoidea [1]

GBenthomangelia aff. obtusangula Reg-mov Pred Epi Based on Conoidea [1;45]

GOenopta kochi Reg-mov Pred Epi Based on Conoidea [1;45]

GSorgenfreispira tenella Reg-mov Pred Epi Based on Conoidea [1;45]

GInquisitor (s.l.) borealis Reg-mov Pred Epi Based on Conoidea [1;21;45]

GRaphitoma spinosoreticulata Reg-mov Pred Epi Based on Conoidea [45]

GTeretia anceps Reg-mov Pred Epi Based on Conoidea [45]

GTeretia sp. Reg-mov Pred Epi Based on Conoidea [45]

GPseudotoma sp. Reg-mov Pred Epi Based on Conoidea [1;18;45]

GGemmula sp. 1 Reg-mov Pred Epi Based on Gemmula [1;18;45]

GGemmula sp. 2 Reg-mov Pred Epi Based on Gemmula [1;18;45]

G‘Gemmula’ sp. 3 Reg-mov Pred Epi Based on Gemmula [1;18;45]

GUnedogemmula (s.l.) hanseata Reg-mov Pred Epi Based on Gemmula [1;18;45]

GActeon sp. Reg-mov Pred Epi Based on Acteon [1;51]

GRingicula promarginata Reg-mov Pred Epi Based on Ringicula [1;14]

GRingicula tiedemanni Reg-mov Pred Epi Based on Ringicula [1;14]

GPyrunculus cf. elongatus Reg-mov Pred Epi Based on other Retusidae [1;42;47]

GVolvulella acuminata Reg-mov Herb Epi Based on Volvulella [1;28]

GCylichna pseudoconvoluta Reg-mov Pred Epi Based on Cylichna [1;28]

GDiaphana moerchi Reg-mov - Epi Based on Diaphanidae [2;31]

GScaphander sp. Reg-mov Pred Epi Based on Scaphander [1;12]

GRoxania sp. Reg-mov Pred Epi Based on Philinoidea [1;2]

GChrysallida sp. Reg-mov Par Epi Based on Pyramidellidae [1;15]

GEulimella cf. acicula Reg-mov Par Epi Based on Pyramidellidae [1;15]

GEulimella cf. scillae Reg-mov Par Epi Based on Pyramidellidae [1;15]

GEulimella sp. 1 Reg-mov Par Epi Based on Pyramidellidae [1;15]

GEulimella sp. 2 Reg-mov Par Epi Based on Pyramidellidae [1;15]

GEulimella sp. 3 Reg-mov Par Epi Based on Pyramidellidae [1;15]

GMegastomia tuexeni Reg-mov Par Epi Based on Pyramidellidae [1;15]

GOdostomia sp. Reg-mov Par Epi Based on Pyramidellidae [1;15]

GParthenina cf. indistincta Reg-mov Par Epi Based on Pyramidellidae [1;15]

(continued)

CAROBENE ET AL.: PALAEOENVIRONMENT OF GROß PAMPAU 7

due to the presence of trends, we detrended the data by

applying generalized differencing (McKinney 1990) with

the R script by Graeme T. Lloyd (http://www.

graemetlloyd.com/pubdata/functions_2.r). All analyses

were computed in R (RStudio Team 2020).

Comparison of benthic molluscan faunas from the North

Sea Basin

To assess the variability of benthic molluscan faunas in

the North Sea Basin that existed penecontemporaneously

with the Groß Pampau fauna, we focused on the Late

Miocene faunal assemblages of the locality of Gram (stra-

totype of the Gram Formation) in southwestern Denmark

that were documented with quantitative data.

In the borehole Gram BH I (Geological Survey of Den-

mark, File No. 141.277), Rasmussen (1966) recognized

five zones defined by the composition of the benthic mol-

luscan assemblages. Applying the regional benthic mol-

luscs zonation, molluscan zones I and II belong to the

Astarte vetula–Aquilofusus luneburgensis (=Langenfeldian

regional stage) Zone, while zones III–V correspond to the

A. reimersi–A. semiglaber (=Gramian regional stage)

Zone (Rasmussen 1966). The correlation of molluscan

zones I and II with the studied section of Groß Pampau

is supported by biostratigraphical evidence, that is, the

occurrence of the index fossil Aquilofusus luneburgensis

and the planktonic gastropods Limacina valvatina,

L. gramensis and L. ingridae (Rasmussen 1966;G

€

urs &

Janssen 2002).

To circumvent taxonomic uncertainties at the species

level, the faunal comparison was conducted at the genus

level, and the original taxonomic assignments in Rasmus-

sen (1966,1968) were updated following Schnetler (2005)

and Wienrich (2007). Given that quantitative data of the

Gram BH I were sampled in a similar way to the data

from Groß Pampau, we selected 20 bulk samples from

zones I–IV that comprise 2373 individuals (54% bivalves,

42% gastropods, 4% scaphopods) belonging to 78 differ-

ent genera (52 gastropods, 23 bivalves, 3 scaphopods).

The final database consists of 50 samples with a total of

9181 individuals belonging to 101 molluscan genera (Car-

obene et al.2023, datasheets 3, 4). Bulk samples with less

than 28 individuals (samples 6 and 7 from Zone I) and

MOLs contributing less than 2% of the overall functional

variability (non-motile semi-infaunal byssate suspension

feeders; non-motile epifaunal pedically attached suspen-

sion feeders) were excluded from the quantitative ana-

lyses. The Q-mode cluster analyses were performed with

the same distance matrix and clustering criteria as applied

in the faunal and functional analyses of the Groß Pampau

benthic fauna. To prevent oversplitting of data, we chose

a lower significance level (p <0.005) for the SIMPROF

test.

Geochemical analysis

To detect potential changes in environmental parameters

during the study interval, we carried out geochemical

analyses of both bulk sediment (concentrations and stable

isotopes of organic carbon and nitrogen) and shells (sta-

ble isotopes of carbon and oxygen). For d

C analysis of

bulk sedimentary organic matter, c. 50 mg of bulk sedi-

ment powder was weighed into a silver foil cup and trea-

ted with 2 molar HCl until bubbling ceased. After drying

the decalcified samples at 40°C, stable isotope analysis

and concentration measurements of nitrogen and carbon

were performed simultaneously with a THERMO/

TABLE 2. (Continued)

Class Taxon Mobility Feeding Tiering Remarks and Source

GTurbonilla cf. koeneniana Reg-mov Par Epi Based on Pyramidellidae [1;15]

GTurbonilla sp. Reg-mov Par Epi Based on Pyramidellidae [1;15]

SFissidentalium floratum Fac-mob(un) Sub-dep Shal-inf Based on Fissidentalium [2;9]

SFissidentalium twistringense Fac-mob(un) Sub-dep Shal-inf Based on Fissidentalium [2;9]

SLaevidentalium sp. Fac-mob(un) Sub-dep Shal-inf Based on Fissidentalium [2;9]

SPolyschides weinbrechti Fac-mob(un) Sub-dep Shal-inf Based on Polyschides [2;47]

Br Cryptopora nysti Non-mot(ped) Susp Epi Based on Cryptopora [8]

†

Numbers in brackets refer to the source of information listed in Appendix S1. B, Bivalvia; Br, Brachiopoda; G, Gastropoda; S,

Scaphopoda. Mobility level: Fac-mob(at), facultative mobile, attached; Fac-mob(un), facultative mobile, unattached; Non-mot(bys),

non-motile, byssate; Non-mot(ped), non-motile, pedically attached; Reg-mov, regularly moving. Tiering: Deep-inf, deep-infaunal; Epi,

epifaunal; Semi-inf, semi-infaunal; Shal-inf, shallow-infaunal. Feeding mechanism: Chem, chemosymbiosis, deposit feeder; Herb, herbi-

vore/grazer; Par, carnivore/parasitic; Pred, carnivore/predator; Sca, carnivore/scavenger; Sub-dep, subsurface deposit feeder; Surf-dep,

surface deposit feeder; Susp, suspension feeder.

8PAPERS IN PALAEONTOLOGY

Finnigan MAT V isotope ratio mass spectrometer,

coupled to a THERMO Flash EA 1112 elemental analyser

via a THERMO/Finnigan Conflo IV interface in the stable

isotope laboratory of the Museum f€

ur Naturkunde, Ber-

lin. Standard deviation for repeated measurements of lab-

oratory standard material (peptone) was <0.15&for both

nitrogen and carbon. Standard deviations of concentra-

tion measurements of replicates of our laboratory stan-

dard are <3% of the concentration analysed. Additionally,

we calculated the molar total organic carbon to nitrogen

ratio (TOC/TN).

We measured the stable carbon and oxygen isotope

compositions of shells of four molluscan species: two

benthic (Gemmula sp.1 and sp.2) and two planktonic gas-

tropods (Limacina valvatina and L. ingridae). Due to their

discontinuous temporal distribution, specimens of Gem-

mula sp.2 and L. valvatina were selected from the lower

part of the section (600–300 cm and 600–200 cm, respec-

tively), while individuals of Gemmula sp.1 and Limacina

ingridae were picked from the remaining part of the sec-

tion. To assess interspecies variations in isotopic composi-

tion and potential vital effects, we analysed respective

congeneric specimens from two samples: S16 for Gem-

mula spp. and S11 for Limacina spp.

The analyses include a total of 31 and 158 well-

preserved specimens of Gemmula spp. and Limacina spp.,

respectively. Owing to their small shell size, on average

five specimens of Limacina spp. were analysed for each

sample. Shells were cleaned with ethanol in an ultrasonic

bath. Entire single shells (Gemmula spp.) and multiple

shells (Limacina spp.) were reduced to a fine powder with

an agate mortar and pestle. Approximately 100–400 lgof

powdered material was put into a clean 10 ml exetainer

and sealed with a septum cap (caps and septa for LABCO

exetainer 438b). The remaining air was removed by flush-

ing the exetainer with He (grade 4.6) for 6 min at a flow

of 100 ml per minute. After flushing, c.30ll of anhy-

drous, phosphoric acid was injected through the septum

into the sealed exetainer using a disposable syringe. After

c. 1.5 h of reaction time at 50°C, the samples were ready

for isotope measurement.

The oxygen and carbon isotope composition in the

in the headspace was measured using a Thermo Fin-

nigan GASBENCH II coupled online with a Thermo Fin-

nigan Delta V isotope ratio mass spectrometer. Reference

gas was pure CO

(laboratory grade 4.5) from a cylinder

calibrated against the VPDB (Vienna PeeDee Belemnite)

standard using IAEA (International Atomic Energy

Agency) reference materials (NBS 18, NBS 19). Reproduc-

ibility of replicate measurements of laboratory standards

(limestone) was <0.10&(one standard deviation). All sta-

ble isotope ratios are expressed in the conventional delta-

notation (d

C, d

O, d

N) in per mil (&) versus AIR

nitrogen and VPDB standard.

RESULTS

Sedimentological analysis

The mean grain size of the sediment samples is 5.3 phi

(range 4.9–5.7 phi). Most samples can be classified as

coarse silt and only a few as very coarse silt. In the

Folk (1954) silt-sand-clay triangle diagram, all samples

are classified as sandy silt (Fig. 3). The mean level of sort-

ing is 1.5 (range, 1.1–1.9), reflecting poor sorting. Many

samples show a bimodal, and some a polymodal grain

size distribution. In the field, distinct primary sedimen-

tary structures were not observed in the studied section,

which suggests mixing of sediment by bioturbators. Trace

fossils occurred throughout the section, except for the

lowermost samples (samples 30–28).

The clay mineral composition of the clay fraction of

the samples consists of illite (mean, 45%; range, 31–

60%), smectite (mean, 31%; range, 21–45%), kaolinite

(mean, 21%; range, 11–30%) and chlorite (mean, 2%;

range, <1–4%). Along the section, clay mineral composi-

tion and the IK/SC ratio vary irregularly with no trend or

cyclicity discernible. A correlation test of clay mineral

content and grain size parameters (mean, sorting, skew-

ness, kurtosis) did not show any significant correlations.

Faunal composition and preservation

The macrobenthic fauna of Groß Pampau (Carobene

et al.2023, datasheet 1) is dominated by bivalves (3619

individuals, 26 taxa) in terms of number of individuals

FIG. 3. Position of the Upper Miocene sediment samples of

Groß Pampau in the silt-sand-clay triangle diagram of

Folk (1954). Abbreviations: C, clay; cS, clayey sand; M, mud; mS,

muddy sand; S, sand; sC, sandy clay; sM, sandy mud; sZ, sandy

silt; zS, silty sand; Z, silt.

CAROBENE ET AL.: PALAEOENVIRONMENT OF GROß PAMPAU 9

and by gastropods (2816 individuals, 81 taxa) in terms of

number of taxa, followed by scaphopods (373 individuals,

4 taxa) and brachiopods (2 individuals, 1 species). Figure 4

shows representative examples of the numerically most

abundant molluscan species considered in this study. With

regard to life habit and mobility, infaunal taxa (63.3% of

the total number of individuals) and regularly moving taxa

(75.4%) are most abundant, while sessile species are

extremely scarce (0.04%). As to feeding mode, the fauna is

composed of deposit feeders (40.8%), carnivores (36.0%)

and suspension feeders (18.4%), with herbivores and che-

mosymbiotic taxa being very rare (4.3%). Except for the

two species of Astarte, which have moderate values for

valve articulation, the percentage of articulated shells of

infaunal bivalves is very high (Table 1). Hence, the high

degree of articulation and frequently preserved micro-

ornamentation on larval and adult shells suggest that

reworking and transport of shells was insignificant, and

shells were deposited in their original habitat.

Faunal associations

Cluster analysis resulted in two groups of faunal assem-

blages separated from each other at a high hierarchical

level, here termed cluster A and cluster B, with the latter

FIG. 4. Illustrations of numerically abundant macrobenthic molluscs from the Upper Miocene of the studied Groß Pampau section.

A, Yoldiella cf. pygmaea.B,Yoldiella spiegleri.C,Astarte gleuei.D,Astarte vetula.E–F, Varicorbula gibba.G–H, Alveinus nitidus.I–J,

Euspira helicina auct. K, Fissidentalium floratum.L,Polyschides weinbrechti.M–N, Aquilofusus luneburgensis.O–P, Bathytoma jugleri.

Q–R, Conilithes poulsenii.S–T, Benthomangelia aff. obtusangula.U–V, Gemmula sp.1. W–X, Gemmula sp.2. Y, Pyrunculus cf. elongatus.

All scale bars represent 1 cm.

10 PAPERS IN PALAEONTOLOGY

being subdivided into five statistically significantly differ-

ent subclusters B1–B5 (Fig. 5). One individual assemblage

(S11), statistically distinct from any other assemblage

according to the similarity profile test, was included in

faunal association B2 upon visual inspection because of

its high similarity with the other assemblages of B2

(Fig. 5; Fig. S1, Table S1). NMDS ordination confirms

the separation of the six faunal associations along NMDS

axis 1 (Fig. 6A). The only overlap in NMDS space occurs

for B2 and B5, and is generated by including the isolated

assemblage S11 in B2.

Comparison of the faunal associations shows that dif-

ferences in the relative abundance of the most common

taxa are only moderate between assemblages, indicating a

fairly even taxonomic composition of the fauna (Fig. 5;

Fig. S1, Table S1). None of the species reaches more than

30% relative abundance in any assemblage. Two small-

sized yoldiid bivalve species, Yoldiella cf. pygmaea and

Yoldiella spiegleri, are the most abundant faunal elements

in all associations (Fig. 5; Table S1). At the highest hierar-

chical level, cluster A differs from B by having relatively

high abundances of the bivalves Varicorbula gibba and

Astarte vetula. In contrast, the gastropod Gemmula sp.1 is

a common constituent of most assemblages in cluster B

(except for B3), whereas it is absent from A. In cluster B,

faunal association B1 has relatively high proportional

abundances of the bivalve Astarte gleuei, whereas this spe-

cies is absent or rare in other assemblages. The bivalve

Alveinus nitidus reaches fairly high abundances in B2, and

the gastropod Gemmula sp.2 is common in faunal associ-

ation B3. Otherwise, although being statistically distinct,

the differences between faunal associations in cluster B

are small, amounting to only minor differences in the

abundance of a few species.

FIG. 5. Heat-map diagram of a two-way hierarchical cluster analysis of Late Miocene faunal assemblages (Q-mode) and benthic

macroinvertebrate species (gastropods, bivalves, scaphopods, and brachiopods) (R-mode) from Groß Pampau based on the relative

abundance of species. Statistically distinct clusters of faunal assemblages S1–S30 define six faunal associations. Colour coding indicates

the relative abundance of species. This figure shows only the segment with species attaining at least 5% relative abundance in any fau-

nal assemblage. The complete two-way cluster diagram, comprising 112 species, is provided in Figure S1.

CAROBENE ET AL.: PALAEOENVIRONMENT OF GROß PAMPAU 11

FIG. 6. Ordination analyses on faunal and functional composition of Late Miocene faunal assemblages from Groß Pampau. A, non-

metric multidimensional scaling ordination (NMDS) using species composition; colour coding and convex-hull outlines reflect Q-

mode clusters of faunal assemblages in Figure 5. B, NMDS of modes of life (MOLs). Colour coding and convex-hull outlines reflect

grouping of assemblages in Figure 7.

12 PAPERS IN PALAEONTOLOGY

MOL associations

We identified 10 distinct modes of life (Fig. 7; Table S2),

of which three are especially abundant: regularly moving

infaunal deposit feeders (MOL1), regularly moving epi-

faunal carnivores (MOL2) and facultatively mobile infau-

nal suspension feeders (MOL3). MOL-based cluster

analysis shows separation of faunal assemblages into two

groups at a high hierarchical level, referred to as MOL

clusters 1 and 2, respectively (Fig. 7). MOL cluster 1

differs from 2 by having particularly high abundances of

infaunal deposit feeders and epifaunal carnivores, whereas

in MOL cluster 2 infaunal suspension feeders constitute

an important additional functional component. At lower

hierarchical levels, each of the two main clusters is subdi-

vided into statistically distinct subgroupings (MOL associ-

ations 1a, b, c and 2a, b). However, differences between

subgroupings in each of the main MOL clusters, while

evident in the ranking of the constituent MOLs, are fairly

minor (Fig. 7; Table S2). In MOL association 1a, infaunal

FIG. 7. Heat-map diagram of a two-way hierarchical cluster analysis of Late Miocene assemblages (Q-mode) and benthic macroin-

vertebrate species (R-mode) from Groß Pampau based on the relative abundances of modes of life (MOLs). Statistically distinct clus-

ters of faunal assemblages S1–S30 define five MOL associations. Colour coding indicates the relative abundance of modes of life. In

MOL8, ‘other feeding’ comprises chemosymbionts and surface deposit feeders.

CAROBENE ET AL.: PALAEOENVIRONMENT OF GROß PAMPAU 13

deposit feeders (Yoldiella spp.) are most abundant and

are followed by epifaunal carnivores, typified by the gas-

tropods Bathytoma jugleri,Benthomangelia aff. obtusan-

gula,Gemmula sp.2 and Pyrunculus cf. elongatus. In MOL

associations 1b and 1c, epifaunal carnivores are dominant,

mainly consisting of Conilithes poulsenii,Bathytoma

jugleri,Benthomangelia aff. obtusangula and Gemmula

spp., whereas infaunal deposit feeders are ranked second.

The relatively high proportions of infaunal suspension

feeders in MOL associations 2a and 2b are partitioned

between several bivalve species, particularly Varicorbula

gibba,Alveinus nitidus,Astarte gleuei and Astarte vetula.

NMDS ordination based on functional groups supports

the separation of MOL clusters 1 and 2 along NMDS axis

1 with some overlap around axis scores of 0.0 (Fig. 6B).

Functional similarity between associations of MOL cluster

1 is also evident from the overlap of the convex-hull

polygon of MOL association 1b with both 1a and 1c. The

arrangement of their faunal assemblages reflects the shift

in dominance from infaunal deposit feeders in 1a to epi-

faunal carnivores in 1b, which is even more pronounced

in 1c. Separation of MOL associations 2a and 2b is gener-

ated by the prevalence of infaunal suspension feeders in

2a, a feature unique to this MOL association.

Temporal patterns

The two dominant Yoldiella species are well represented

throughout the succession, whereas distinct temporal

changes in relative abundance between common species

are observed in four bivalve species (Astarte gleuei,Astarte

vetula,Alvenius nitidus and Varicorbula gibba) and two

gastropod taxa (Gemmula sp.1 and Gemmula sp.2)

(Fig. 8). Alvenius nitidus and Varicorbula gibba show sim-

ilar temporal abundance fluctuations, with relatively high

values in the lower part of the succession (c. 600–440 cm

height) and again from c. 240 to 140 cm. Astarte vetula

and Astarte gleuei occur in the lower and the upper part

of the section, respectively. Likewise, Gemmula sp.2

becomes rare in the upper half of the section, where

Gemmula sp.1 is prominent.

The temporal distribution of the six faunal associations

exhibits a pattern of general stability in faunal composi-

tion across several consecutive assemblages before shifting

to a new faunal configuration (Fig. 9). Recurrence of a

faunal association at different levels of the succession is

evident only in faunal association B5, occurring from 280

to 220 cm and again from 140 to 100 cm. Given the hier-

archical nature of clusters, the main shift in species

FIG. 8. Relative abundances of the most common Late Miocene bivalve and gastropod species at Groß Pampau.

14 PAPERS IN PALAEONTOLOGY

composition and abundance is from faunal association A

to cluster B and occurs only once: from S22 to S21 at

420 cm.

Assemblage-level rarefied richness fluctuates moderately

in a narrow band of 20–30 species without any distinct

temporal trend (Fig. 10). Richness appears to fluctuate

somewhat more in the lower third of the succession com-

pared with younger parts (Fig. 10). Similarly, Simpson’s

index is very stable and indicates fairly consistent, very

even species abundance distributions.

Temporal changes in the relative abundance of MOLs

are apparent in regularly moving infaunal deposit feeders

(MOL1), regularly moving epifaunal carnivores (MOL2)

and facultatively mobile infaunal suspension feeders

(MOL3). The increases in relative abundance of MOL1

and MOL2 in the lower third of the section correspond

to a decrease of MOL3 (Fig. 11A). Thereafter, the per-

centage of MOL1 remains fairly stable whereas MOL2

und MOL3 exhibit a longer-term undulation in opposing

directions. Considering the three main ecological catego-

ries (mobility level, tiering, and feeding mechanism) sepa-

rately, we noticed that changes in feeding habits account

for most of the MOLs’ temporal variability (Fig. 11B).

Deposit feeders gradually increase in the lower third of

the section, becoming relatively abundant and stable

thereafter. Carnivores show a similar increasing trend, but

fluctuate considerably in relative abundance in the upper

part of the section. Suspension feeders are abundant in

the lowermost samples but then decline across the lower

third of the section, stay fairly stable in the middle third,

and slightly fluctuate in the upper third of the section.

The temporal distribution pattern of the MOL associa-

tions resembles that of the faunal associations, suggesting

ecological stability punctuated by shifts to a different

functional composition that again remains stable for sev-

eral consecutive assemblages (Fig. 9). According to cluster

hierarchy, the main shift in species composition and

abundance from faunal association A to cluster B occurs

simultaneously with an ecological shift from MOL cluster

2 to cluster 1 at 420 cm. Also, most other shifts among

associations B1–B5 occur contemporaneously or in close

temporal proximity with shifts of MOL associations. Sim-

ilar to taxonomic diversity, functional diversity indices

are also fairly constant with no long-term trend and with-

out any abrupt shifts anywhere in the studied interval

(Fig. 10).

Comparison of molluscan associations of Groß Pampau and

Gram, Denmark

Using genus composition, the Q-mode cluster analysis

combining faunal samples from Groß Pampau and Gram

resulted in 11 faunal associations (Fig. 12A; Table S3).

The associations for Gram mirror the molluscan assem-

blages already identified by Rasmussen (1966). Cluster

analysis shows a complete separation between the benthic

molluscan assemblages of Gram and Groß Pampau. This

pattern also holds when the clustering is based on

presence–absence data (not shown). The bivalve Yoldiella

dominates most faunal associations at both sites, except

for Gram associations 1 and 6 (Table S3). In addition,

quantitatively important faunal elements in Gram are the

bivalve genera Limopsis,Cyclocardia,Astarte and Goodal-

lia, and the gastropods Tritia and Obtusella, while in

Groß Pampau the faunal associations are characterized by

the numerically abundant bivalve genera Astarte,Varicor-

bula,Alveinus and a few prominent gastropod genera

(Table S3).

In terms of ecological composition, the MOL-based

cluster analysis recognized two main clusters, A and B,

the latter being composed of three subclusters, labelled

MOL associations B1, B2, and B3 (Fig. 12B). In all four

MOL associations the same three modes of life prevail

albeit in variable relative abundances (Table S4). At the

highest hierarchical level, cluster A is unique in being

dominated by infaunal suspension feeders, whereas in

B1–B3 infaunal deposit feeders and epifaunal carnivores

FIG. 9. Stratigraphical distribution of Late Miocene benthic

macroinvertebrate associations at Groß Pampau based on species

composition (left) and their modes of life (MOLs, right). See

cluster analyses in Figures 5and 7, and Tables S1 and S2, for

assignment of faunal assemblages to associations.

CAROBENE ET AL.: PALAEOENVIRONMENT OF GROß PAMPAU 15

occupy first or second rank. As evident from the assem-

blages assigned to MOL associations A, B2 and B3, the

functional variability of Gram associations closely reflects

the one recorded in the lower third of the Groß Pampau

section. MOL associations A and B2, which include most

of the Gram samples (73% and 85% of each association,

respectively), are dominated by infaunal suspension

feeders (39.1%) and epifaunal carnivores (33.4%), respec-

tively. In contrast, MOL association B3, mainly repre-

sented by Groß Pampau samples (70%), yields a high

percentage of infaunal deposit feeders (33.6%) and similar

abundances of epifaunal carnivores (25.9%) and infaunal

suspension feeders (21.8%). MOL association B1, solely

consisting of Groß Pampau samples, is dominated by epi-

faunal carnivores (39.7%) and infaunal deposit feeders

(37.5%).

Stable carbon and nitrogen isotope signatures of bulk

sediment, TOC and TOC/TN

The concentration of TOC and TN, the TOC/TN ratio,

and the d

C and d

N stable isotope composition of bulk

sediment are shown in Figure 13 and listed in Carobene

et al.(2023, datasheet 5). All geochemical profiles are rel-

atively stable and consistent over time (Fig. 13). Nitrogen

varies between 0.06% and 0.08%, with an average of

0.07 0.01%, while TOC ranges from 0.92% to 1.76%,

with a mean of 1.26 0.23%. The TOC/TN ratio varies

from 11.39 to 23.95, with a mean of 15.95 2.90

(Fig. 13). The d

C composition of organic carbon varies

between 24.61&and 23.48&, with a mean of

24.03 0.32&. The d

N signature of organic matter

ranges from 3.05&to 5.31&, with a mean of

4.19 0.44&.

Stable carbon and oxygen isotope signatures of benthic and

planktonic gastropod species

The stable isotope compositions of both benthic and

planktonic gastropod species are given in Figure 14 and

Carobene et al.(2023, datasheets 6, 7). Except for two

outliers (samples S16 and S26), the d

O composition of

Gemmula sp.1 and sp.2 is relatively similar and constant

throughout the section (G. sp.1: mean, 1.82 0.14&;G.

sp.2: mean, 1.80 0.62&). The outliers of Gemmula sp.2

in S26 and S16 are most likely to represent false measure-

ments rather than true signals of environmental change

(S26) or species-specific vital effects (S16). In the lower

part of the section, the d

C composition of Gemmula

sp.2 ranges from 0.36&to 1.56&with two outliers (S30

and S27), while in the upper part the d

C values of Gem-

mula sp.1 remain more stable (0.10–1.04&). Overall, the

FIG. 10. Macrobenthic abundance and diversity through the Upper Miocene succession of Groß Pampau. Absolute abundance

(number of individuals per constant amount of sediment) and various biodiversity metrics of Late Miocene molluscan assemblages

based on their taxonomic and ecological composition. Grey lines represent the three-point moving average.

16 PAPERS IN PALAEONTOLOGY

C profile shows a gradual decrease, followed by a slight

increase towards the top of the section.

No measurements could be obtained for L. ingridae in

S4, while the isotopic composition of specimens from

samples 1, 2 (L. ingridae), 21, 23 and 25 (L. valvatina)

were not considered further because sample sizes were

inadequate for isotope analyses. The d

O values of

L. valvatina and L. ingridae display a slightly greater vari-

ability than those of the benthic Gemmula species with

values ranging from 1.84&to 1.32&and from

2.41&to 0.50&, respectively. Except for outliers in

samples 17, 12 and 11, the stable oxygen values of both

species are fairly similar, suggesting the lack of species-

specific isotopic fractionation. This is further supported

by the concomitantly low d

O value of L. valvatina in

S11, indicating an environmental change as the source of

the signal. The composite d

O profile exhibits a fluctuat-

ing pattern shortly interrupted by a negative shift at 2.1–

1.9 m. The d

C composition of Limacina species records

a single outlier in S11 and a fluctuating pattern with

mean values of 1.82 0.31&for L. valvatina and

1.57 0.50&for L. ingridae.

Palaeotemperature reconstructions

For the interval from the Middle–Late Miocene boundary

into the early Late Miocene, the d

O values of seawater

are estimated to range between 0.5&and 0.25&stan-

dard mean ocean water (SMOW) (Lear et al.2000; Bill-

ups & Schrag 2002). Palaeotemperature estimates from

the d

O composition of benthic and planktonic species,

derived from the equation of B€

ohm et al.(2000), along

with estimated d

O values of seawater, are presented in

Figure 14 and Carobene et al.(2023, datasheets 6, 7).

While inferred bottom-water temperatures remain rela-

tively stable throughout the succession, sea surface tem-

peratures oscillate slightly around the mean, with a

transient increase in samples 11 and 10. Except for these

two samples and disregarding the dubious d

O values of

Gemmula in S26 and S16 (see above), the bottom and

surface temperatures calculated from the remaining sam-

ples (and assuming a d

seawater

value of 0.5&

SMOW), have mean values of 9.7 0.7°C and

18.7 2.6°C, respectively. When using a d

seawater

0.25&SMOW, palaeotemperature estimates simply

FIG. 11. Abundance of macrobenthic functional groups through the Upper Miocene succession of Groß Pampau. A, relative abun-

dance (%) of the most important modes of life (MOLs). B, relative abundance of the major ecological parameters: feeding mechanism

(deposit feeders, carnivores, suspension feeders, predators), mobility level (regularly moving) and tiering (infaunal). Grey lines repre-

sent the three-point moving average. Abbreviations: n, number of species; N, number of individuals; MOL1, regularly moving, infaunal,

deposit feeder; MOL2, regularly moving, epifaunal, carnivore; MOL3, facultatively mobile, infaunal, suspension feeder. For less abun-

dant MOLs see Figure 7.

CAROBENE ET AL.: PALAEOENVIRONMENT OF GROß PAMPAU 17

FIG. 12. Q-mode cluster analyses based on the genus-level composition of quantitative samples of benthic macroinvertebrates from

Groß Pampau and Gram (Denmark) considering the relative abundances of: A, species; B, ecological groups. Sample coding: GB, Gram

borehole I; GB–I, Gram borehole, Zone I; GB–II, Gram borehole, Zone II; GB–III, Gram borehole, Zone III; GB–IV, Gram borehole,

Zone IV; GP, Groß Pampau. Zones numbered after Rasmussen (1966).

18 PAPERS IN PALAEONTOLOGY

increase by 1.1°C. The offset between surface- and

bottom-water temperatures shows a mean value of

8.8 2.6°C.

Canonical correspondence analysis and Spearman’s rank-

order correlation

CCA axis 1 (CCA1) and CCA axis 2 (CCA2) explain

31.3% of the total variance in the observed species abun-

dances (total inertia =0.64) and 76.9% of the variance in

weighted averages and class total of species with respect

to the environmental variables (Fig. 15; Table S5). As

shown in Table 3, assemblage structure is significantly

associated with TOC (CCA dimension 1, biplot

score =0.97, Eigenvalue =0.14, F-value =8.14, p =0.001,

Table S5) and marginally influenced by TN content (CCA

dimension 2, biplot score =0.72, Eigenvalue =0.02, F-

value =3.16, p <0.1, Table S5). In contrast, neither the

grain size and clay mineral ratio of the sediment nor the

stable isotope ratios of nitrogen and of organic carbon

were significantly correlated with assemblage patterns.

Taxa and faunal samples clustering at the centre of the

ordination are either not correlated to any of the environ-

mental variables or occur throughout the section, as in

the case of Yoldiella spiegleri and Y. pygmaea (Fig. 15). In

contrast to Gemmula sp.2 and A. vetula,Gemmula sp.1

and A. gleuei were abundant at the higher TOC values in

the upper part of our studied section. Alveinus nitidus

and R. tiedemanni are found at intermediate positive

CCA2 values, while V. gibba plots at negative CCA1

scores.

When analysing the temporal distributions of the abun-

dant species separately, Spearman’s rank-order correla-

tions indicated no significant correlations of Y. spiegleri,

Y. cf. pygmaea,A. vetula, the two species of Gemmula,

and F. floratum with any of the analysed environmental

factors (Table S6). A significant positive correlation was

found between the distribution of A. nitidus and d

(q=0.46, p =0.01), and V. gibba and the TN content

(q=0.37, p =0.05), and a negative correlation exists

between A. gleuei and the clay mineral ratio (q=0.38,

p=0.05). Thus, with the exception of V. gibba, no other

species was significantly linked to the variables that domi-

nated assemblage variation (i.e. TOC and TN). However,

assemblage structure showed variation along these

FIG. 13. Geochemical and sedimentological data of Upper Miocene bulk rock samples from the Groß Pampau section. Shown are

relative abundances of total organic carbon (TOC) and nitrogen (TN), their isotopic signatures (d

C, d

N), the molar total organic

carbon to total nitrogen ratio (TOC/TN), the clay mineral ratio (IK =illite +kaolinite; SC =smectite +chlorite), and the mean grain

size distribution. Grey lines represent the three-point moving average.

CAROBENE ET AL.: PALAEOENVIRONMENT OF GROß PAMPAU 19

environmental gradients (Fig. 15). The earliest faunal

assemblages S30–S28 are associated with relatively low

TOC and high TN values. Faunal assemblages S27–S24

correspond to a fall in TN to lower values, which is con-

tinued for assemblages S23–S18. Assemblages S12–S7 are

found at higher TN (and d

C) values, while S4–S1 are

associated with the highest TOC values. Although these

changes are not large, they correspond to the subtle tem-

poral changes in TN and TOC, showing that assemblages

and environmental conditions are slowly drifting in con-

cert over time.

DISCUSSION

Reconstruction of the Late Miocene benthic ecosystem of

Groß Pampau

Our combined sedimentological, geochemical and faunal

evidence allows for a reconstruction of the environmental

conditions during deposition of the Groß Pampau succes-

sion. Overall, the relatively monotonous sedimentary suc-

cession, coupled with low variability in geochemical

proxy data and relatively high and fairly stable values of

both species richness and evenness, point to fairly uni-

form environmental conditions, which were generally

favourable for molluscan communities. In the following,

we reconstruct the Groß Pampau ecosystem in more

detail in terms of depositional processes, water depth and

water energy, oxygenation, substrate conditions, water

temperature and productivity.

Depositional processes and sediment sources. Poor sorting

and the often bimodal, partly polymodal grain size distri-

bution suggest differently transported sediment fractions.

Although clay and silt were likely to have been deposited

by the background settling of suspension load, the sand

fraction of the sandy silts might be transported by events

such as storms or turbidity currents.

The clay mineral composition of the sediment samples

reflects different source areas as well as a climatic influ-

ence. Illite, which dominates the clay fractions of most

samples, is most likely to have originated predominantly

from physical weathering of gneisses and granites of the

Precambrian Fennoscandian Shield and was transported

to the North Sea by the west-flowing Eridanos River

(Gibbard & Lewin 2016). Illite formation was probably

favoured by the relatively cooler climate in the Late

FIG. 14. Carbon and oxygen isotopic composition of the shells of benthic (Gemmula sp.1 and Gemmula sp.2) and planktonic gastro-

pods (Limacina valvatina and L. ingridae) from the Upper Miocene of Groß Pampau. Palaeotemperature estimates (T) are calculated

from the d

O composition of benthic and planktonic species for two different d

O values of seawater (d

seawater

=0.5&and

seawater

=0.25&), and from the absolute temperature, which was computed using the oxygen isotope thermometer of B€

ohm

et al.(2000). Grey lines represent the three-point moving average. Abbreviation: VPDB, Vienna PeeDee Belemnite.

20 PAPERS IN PALAEONTOLOGY

Miocene (Nielsen et al.2015). The Neogene uplift of the

Rhenish Slate Mountains and the Carpathians could have

created another source of illite (Nielsen et al.2015). It is

most likely that smectite was formed primarily by

onshore weathering of Palaeogene volcanic ash derived

from pyroclastic activity associated with the opening of

the North Atlantic (Huggett & Knox 2006) or by its hal-

myrolysis in seawater (Nielsen et al.2015). Kaolinite

probably originated mainly from the chemical weathering

of the Fennoscandian Shield and, as a relatively coarse

clay mineral, was preferentially deposited near the coast

(Biscaye 1965). Chlorite originated mainly from physical

erosion of the Caledonides on both sides of the northern

North Sea (Nielsen et al.2015). Being susceptible to

chemical weathering, its occurrence was most likely to

have been promoted by the cooling of the Late Miocene

FIG. 15. Results of the canonical correspondence analysis in a species-conditional triplot showing the distribution of Late Miocene

assemblages (circles) and species (crosses) from Groß Pampau along the environmental gradients (blue arrows). Species abbreviations:

Ap, Aporrhais sp.; Bo, Benthomangelia aff. obtusangula; Ci, Cingula inusitata auct.; Cg, Colus gregarius; Cp, Conilithes poulsenii; Eh,

Euspira helicina auct; Ff, Fissidentalium floratum; G. sp.1, Gemmula sp.1; G. sp.2, Gemmula sp.2; Ge, ‘Gemmula’ sp.3; Lr, Latirulus

rothi; Ls, Lyrotyphis cf. sejunctus; Ng, Nucula georgiana; Og, Obtusella gottscheana; Pf, Pseudobabylonella cf. fusiformis; Ps, Parvicardium

cf. scabrum; Pw, Polyschides weinbrechti; Rm, Ringicula promarginata; Ta, Teretia anceps; Tm, Tritia mothsi; Tt, Turritellinella tricarinata

auct.; Yg, Yoldia cf. glaberrima; Yp, Yoldiella cf. pygmaea; Ys, Yoldiella spiegleri.Environmental gradient abbreviations:d

org

, organic

carbon isotope composition; d

N, nitrogen isotope composition; GS, mean grain size; IK/SC, clay mineral ratio; TN, total nitrogen

content; TOC, total organic carbon content.

TABLE 3. Biplot scores of environmental variables against CCA dimensions 1 (CCA1) and 2 (CCA2), followed by the significance of

the environmental variables tested using ANOVA-like permutation and the setting ‘by margin’.

Environmental variables CCA1 CCA2 d.f. Chi-squared F-value p-value

TOC 0.97 0.07 1 0.08412 4.868 0.001

TN 0.21 0.72 1 0.03464 2.004 0.077

N0.33 0.28 1 0.02463 1.425 0.184

org

0.21 0.62 1 0.02348 1.359 0.205

IK/SC 0.22 0.25 1 0.01319 0.763 0.595

Mean grain size 0.33 0.04 1 0.01721 0.996 0.409

Residual 22 0.38018

org

, stable carbon isotope composition of organic matter; d

N, stable nitrogen isotope composition; IK/SC, clay mineral ratio

(illite +kaolinite/smectite +chlorite); Mean grain size, mean grain size distribution; TN, total nitrogen content; TOC, total organic

carbon content.

CAROBENE ET AL.: PALAEOENVIRONMENT OF GROß PAMPAU 21

climate. The clay mineral composition of the samples

roughly corresponds to the clay mineral composition of

sediments from offshore areas of the northern North Sea

Basin in the Late Miocene (Nielsen et al.2015). This

indicates the dominance of supra-regional sediment

sources and an offshore depositional environment for the

Groß Pampau sediments.

Water depth and energy. Assessment of the water depth of

the Groß Pampau benthic environment can be obtained

via sedimentological evidence; the proportion of organisms

depending on photosynthetic plants as a food source; and

by the preferred habitat depth of species that still exist

today or that of closely related living species. The monoto-

nous silty facies of the succession indicates that deposition

occurred in a low energy environment, mostly below storm

wave base. Only the occasional very thin layers (5 mm

thick) of very fine-grained sand and thin shell accumula-

tions at the base of the section indicate possible proximity

to storm wave base and are interpreted as distal storm

beds. Low energy conditions mostly below storm wave base

are supported by the high percentage of deposit feeders

suggesting that calm water conditions enabled the accumu-

lation of particulate organic matter at the sea floor. Fur-

thermore, the high percentages of articulated bivalves

corroborate the calm bottom conditions and the general

lack of reworking. The scarcity of herbivores (eight taxa,

4.3% of the total number of individuals) suggests a habitat

at the limit of or beyond the photic zone.

Accurate bathymetric preferences are available only for

the extant species Varicorbula gibba, which commonly

thrives on muddy sand bottoms at 10–40 m depth, and

occasionally reaches greater depths (down to 250 m; Hrs-

Brenko 2006). The living species Yoldiella philippina

(Nyst), morphologically similar to Yoldiella cf. pygmaea,

usually lives at depths of 100–300 m and occasionally has

been found in shallower habitats up to 25 m

(War

en 1989). The preferred habitat depths of the extinct

Astarte vetula and Astarte gleuei are unknown but living

astartids usually thrive at depths of 10–150 m. Alveinus

nitidus is an extinct species of the bivalve family Kellielli-

dae, the representatives of which inhabit a wide range of

habitats, from the continental shelf down to 1000 m

depth (Krylova et al.2018). Gemmula species are also

ubiquitous inhabitants of rather deep environments, rang-

ing from 50 to 500 m depth (Heralde et al.2010). The

bathymetric preferences of common Miocene species at

Groß Pampau are thus consistent with a water depth of

several tens of metres. This estimation agrees with the

minimum depth of 40–60 m inferred by Moths (1994)

based on the occurrence of elasmobranch species.

Oxygenation of bottom waters and the upper sediment

layer. The dominance of shallow infauna and high

diversity and abundances of epifaunal carnivores indicate

that oxygenated conditions prevailed at the sediment–

water interface and in the uppermost layers of the

sediment. However, increased abundances of the opportu-

nistic bivalve Varicorbula gibba in the lower third of the

section hint at intermittently less favourable conditions,

possibly related to transient hypoxia (see below). A posi-

tion of the redox boundary at relatively shallow depth

might explain the absence of deep infaunal burrowers.

Substrate conditions. The burrowing activity of the abun-

dant deposit-feeding nuculanids (Yoldiella spp.) probably

resulted in a moderately soft substrate and a turbid

boundary layer. The fairly soft and potentially turbid bot-

tom conditions and the lack of adequate hard parts may

have prevented the settlement and growth of a diverse

sessile suspension-feeding epifauna (trophic group amens-

alism, Rhoads & Young 1970). Likewise, a soft substrate

and poor penetration of daylight, as suggested by the

scarcity of herbivorous gastropods, may have hampered

the growth of macrophytes and algal mats. Nevertheless, a

soupy consistency of the substrate is unlikely because

morphological adaptations, such as a snowshoe strategy

among epifaunal gastropods, are missing, and many

mobile epifaunal species were able to colonize the sea

floor.

Sea surface and bottom water palaeotemperatures. Stable

isotope composition of molluscan shells is a reliable

proxy for palaeoenvironmental reconstruction (Kobashi &

Grossmann 2003;L



ecuyer et al.2004; Latal et al.2006a,

2006b; Reich et al.2015). Given that secretion of the cal-

careous shell by molluscs usually occurs over several years

and in isotopic equilibrium with the ambient water, their

shells are likely to record changes in the isotopic compo-

sition of ambient water due to global and local palaeoen-

vironmental factors (Grossman & Ku 1986; Wefer &

Berger 1991). The d

O of marine mollusc shells is related

to both the temperature and d

O of the seawater (Leng

& Lewis 2016). Rather than an increase in temperature,

the shift toward isotopically lighter d

O values of Lima-

cina spp. at 2.1–1.9 m is more likely to indicate a change

in surface water salinity, while d

O values in benthic

Gemmula suggest that bottom water salinity (and temper-

ature) remained constant. Although freshwater exhibits a

wide range of isotopic composition, their values are

clearly lower than in marine settings (Latal et al.2006b).

A temporary increase in the discharge of freshwater from

land (e.g. by a shifting river mouth) is expected to lower

the oxygen isotope ratio of surface water, and would

explain the isotopically lower d

O values of Limacina

spp.

With the sampling technique adopted here, the d

values represent averaged isotopic compositions recorded

22 PAPERS IN PALAEONTOLOGY

throughout the growth of a single shell (Gemmula spp.)

or multiple shells (Limacina spp.). Thus, the conversion

of d

O values into seawater temperatures provides corre-

sponding estimates of multi-annual temperatures of both

the mixed surface layer and bottom waters, averaged over

the growth season. The calculated sea surface tempera-

tures of 18.7°C and 19.8°C, depending on the SMOW

values used, are consistent with the continental mean

annual temperature of c.17–19°C estimated for the Mid-

dle to early Late Miocene of northern Europe using paly-

nological proxies (Larsson et al.2011) and correspond to

warm temperate climatic conditions. The pronounced dif-

ference in temperature of c. 8.8°C between the mixed sur-

face layer and bottom waters implies the presence of a

strong seasonal thermocline. An example of a similar

present-day oceanographic situation seems to be the Gulf

of Lions (France) in the western Mediterranean Sea. Here,

a summer thermocline forms at 10–20 m water depth

and separates the bottom water with a minimum temper-

ature of 13.5°C from the surface water with a mean tem-

perature of 20°C (Millot 1990; Requena et al.2013). It

should be kept in mind that, owing to the geochemical

sampling of whole shells, the estimated sea surface tem-

peratures result from a mixed signal of temperature

values realized during the whole period of shell growth.

Accordingly, surface temperatures during the summer

months were probably higher than the calculated average

temperatures, and winter temperatures were lower. Con-

sequently, also the temperature differences between sur-

face and bottom waters represent mixed signals. It is

quite plausible that a strong thermocline with a marked

temperature gradient developed during summer, while

ocean mixing occurred during winter, and surface and

bottom water temperatures converged.

Biogeochemical cycles and trophic conditions. Although the

N of sinking organic nitrogen may be altered by con-

tamination with low-d

N bacterial biomass and by diage-

netic processes, bulk sedimentary d

N is often used for

understanding past and present changes in the ocean

cycle of nitrogen (Galbraith et al.2008; Robinson

et al.2012). The stability of the bulk d

N signal through-

out the studied section excludes any diagenetic alteration.

The d

N signature and TOC concentrations indicate

that mesotrophic open marine conditions and a relatively

well oxygenated sea-floor environment prevailed during

the deposition of the studied succession (Struck 2012).

N values between 3&and 8&and relatively low TOC

concentrations are usually associated with a nitrogen cycle

dominated by nitrate production, in which primary pro-

ducers are provided with nutrients, influencing the flux

of organic matter to the sea floor. The high abundances

of deposit feeders in the macrobenthic assemblages and

the frequent findings of baleen whales point towards a

relatively high productivity rate, sustaining a diverse food

web that also includes carnivores, suspension-feeders and

a few herbivores. An indicator of constant primary pro-

duction is provided by the offset in d

C composition

between the shells of benthic and planktonic species. Ben-

thic Gemmula species have on average a c.1&more

depleted d

C composition than the planktonic Limacina

species, suggesting an isotopic offset between surface and

bottom waters. The d

C of the surface water dissolved

inorganic carbon was probably

C enriched, resulting

from

C assimilation by photosynthetic organisms, while

bottom waters were

C depleted due to decomposition

of isotopically lighter organic matter on the sea floor

(Kobashi & Grossmann 2003).

The sedimentary organic d

C composition and the

TOC/TN ratio accurately reflect the source of organic

matter preserved in coastal environments (Twichell

et al.2002; Mackie et al.2005; Lamb et al.2006). The

C of marine particulate organic carbon and marine

algae is higher compared with that of terrestrial particu-

late organic carbon (Fig. 16; Lamb et al.2006 and refer-

ences therein). When analysed together, the relatively low

C composition of the organic carbon and the TOC/TN

of the samples suggest that the organic matter mainly

consisted of marine organic carbon with a partial contri-

bution of land-derived material (Fig. 16).

Abiotic drivers of faunal variability

Canonical correspondence analysis combines quantitative

species abundance data and environmental data at sites to

obtain environmental gradients that maximize the niche

separation among species (Ter Braak 1986; Ter Braak &

Verdonschot 1995). Our CCA results reveal the influence

of two abiotic environmental variables, TOC content and,

less distinctly, TN content, on the distribution of the

Groß Pampau molluscan fauna (Fig. 15, Table 3). As

proxies of productivity and terrestrial runoff, TOC and

TN are closely related to each other and reflect changes

in organic matter availability at the sea floor, which can

affect the composition and ecological structure of macro-

benthic communities. Notably, as geochemical conditions

remained fairly stable across the section, the TOC and

TN gradients in the CCA represent narrow ranges of vari-

ation. Nevertheless, this slight variation may have chan-

ged the balance of assemblage membership over time in

an ecologically meaningful way, such as by altering com-

petition dynamics.

The clustering of the most common species in the mid-

dle of the ordination plot indicates the similarity in core

members of the Groß Pampau benthic molluscan fauna

and its probable ecosystem functioning over time

(Fig. 15). The distribution of faunal assemblages along

CAROBENE ET AL.: PALAEOENVIRONMENT OF GROß PAMPAU 23

the TOC gradient does not correspond to any substantial

increase in functional groups usually related to organic

enriched sediments, such as deposit feeders or opportu-

nistic species, and is most likely to reflect the generally

low variability in total organic matter content. The posi-

tion of a few species (i.e. Astarte vetula,Gemmula sp.2,

Gemmula sp.1 and Astarte gleuei) along CCA1 suggests

slightly different preferences for organic carbon concen-

trations that are indicative of subtle but statistically signif-

icant assemblage change.

As for CCA2 and the corresponding TN gradient, sev-

eral species, namely Ringicula tiedemanni,Alveinus nitidus

and Varicorbula gibba, were found at intermediate posi-

tive CCA2 axis values and low to negative CCA1 values

(Fig. 15). Thus, part of the faunal assemblages in faunal

association A and those belonging to faunal association

B2, in which these species are common constituents,

apparently co-vary with the small-scale variation in nitro-

gen content. The Spearman correlation test yields only

one positive and significant covariance: between V. gibba

and nitrogen concentration, which is consistent with the

opportunistic character of this species. The dominance of

V. gibba in both recent and fossil communities is usually

associated with unfavourable environmental conditions

and re-colonization of devastated areas (Bernasconi &

Robba 1993; Dominici 2001; Mandic & Harzhauser 2003;

Dominici & Kowalke 2007; Zuschin et al.2007; Schnei-

der 2008; Toma

sovych et al.2018). This species tolerates

a wide range of environmental disturbances: seasonal oxy-

gen depletion, chemical pollutants and increased turbidity

(Yonge 1946; Hrs-Brenko 2006). In Groß Pampau, the

individual abundances of up to c. 20% of V. gibba in the

lower part of the section (faunal association A) coincide

with slightly higher and fluctuating TN proportions

(Fig. 13). Nevertheless, both taxonomic (richness and

evenness) and functional diversity in this part of the sec-

tion show variable but relatively high values (Fig. 10).

Thus, unlike the monospecific assemblages described from

stressful environments, the proliferation and distribution

of V. gibba were probably controlled by short-term, or

low-level environmental stress, such as transient hypoxia

or elevated turbidity. Regarding the other concurrent spe-

cies, low-oxygen tolerance was detected in some extant

species of astartids and ringiculids (Diaz & Rosen-

berg 1995; Dominici 2001). Hence, the co-occurrence of

V. gibba with A. vetula, ringiculids and the generalist

Alveinus nitidus (Schneider 2008) in faunal association A,

and of V. gibba with A. nitidus and Ringicula tiedemanni

in faunal association B2, supports the occasional occur-

rence of less favourable bottom conditions. The positive

correlation of A. nitidus with the d

O composition of the

benthic gastropods and the negative correlation of

A. gleuei with the clay mineral ratio cannot be explained

in terms of ecological requirements of the species. Given

the low variation in d

O composition of Gemmula spp.

and the lack of any significant correlation between the

clay mineral ratio and the temporal distribution of faunal

assemblages, we consider these correlations as spurious.

Punctuated stasis of faunal associations

The temporal distribution of faunal associations describes

a pattern of punctuated stasis, characterized by temporal

persistence in taxonomic and functional composition,

interrupted by occasional shifts to a different species and

MOL composition (Fig. 9). Both external environmental

factors and internal ecological interactions have been pro-

posed to underlie the patterns of stability and shifts in

community structure over time (Miller 1986,1997; Brett

& Baird 1995; Brett et al.1996; Ivany 1996; Jablonski &

Sepkoski 1996; DiMichele et al.2004; Ivany et al.2009).

The stabilizing effect that constant ecological conditions

exert on communities derives from the observation that

taxa with similar environmental preferences tend to

occupy the same habitat for as long as that habitat per-

sists (Ivany 1996 and references therein). Thus, it is very

likely that the stable physical environmental conditions

that occurred throughout the studied succession, as

shown by the sedimentological and geochemical data,

FIG. 16. Organic carbon isotopic composition (d

org

)vs

molar total organic carbon to total nitrogen ratio (TOC/TN) for

the bulk rock samples of Groß Pampau, showing the partial

contribution of land-derived material in the total organic matter

of Groß Pampau. Ranges of d

C and TOC/TN compiled after

Lamb et al.(2006). Abbreviations: DOC, dissolved organic car-

bon; POC, particulate organic carbon.

24 PAPERS IN PALAEONTOLOGY

contributed to the pattern of species consistency in faunal

associations.

As major drivers of community structure, biotic inter-

actions are important in promoting temporal stability of

communities (Connell & Slatyer 1977; Morris et al.1995;

Ivany 1996; Miller 1997). According to recent studies, it

is the diversity of interaction types that determines the

ecological structure of communities in nature (Mougi &

Kondoh 2012; Quian & Akcay 2020). The effects of pre-

dation and competition on community organization have

received the majority of attention (Paine 1966; Menge &

Sutherland 1976; Pearson & Rosenberg 1987; Chase

et al.2002; Holt 2009). Predation can positively or nega-

tively affect community assembly by either enhancing or

hampering competition within the same guilds of prey

(Bodini 1991; Chase et al.2002; Holt 2009). Community

stability and species diversity usually increase when preda-

tors promote the coexistence of competing prey species

(Chase et al.2002 and references therein). For example,

predation on competing prey can alter the diversity and

abundance of the prey’s resources and increase the proba-

bility of coexistence by supporting differences in resource

consumption (Chase et al.2002).

In Groß Pampau, the overall functional structure of

the fauna is characterized by a great diversity and abun-

dance of carnivorous gastropods. These include predatory

species of the shell-drilling families Naticidae (Euspira)

and Muricidae (Lyrotyphis,Murex and Pagodula), which

on average reach 4.7% relative abundance in faunal

assemblages. The importance of successful drilling preda-

tion can be assessed by the frequency of drillholes. We

recorded the frequency of individuals bearing circular

boreholes of millimetric size as typically produced by

naticids and muricids (Fig. 4and Carobene et al.2023,

datasheet 8). The most abundant species at Groß Pam-

pau, the two species of Yoldiella, exhibit very low drilling

frequencies (Carobene et al.2023, datasheet 8), although

their mobile and infaunal lifestyle itself can be seen as an

antipredator strategy (Vermeij 1987; Aberhan et al.2006).

Similarly, drilling frequencies for most other species with

at least one bored specimen are mostly low (less than

3%). Among the highest drilling frequencies are those

found in the two abundant species of Astarte (8.7% and

8.1% on average, respectively). An even higher frequency

is associated with Turritellinella tricarinata (17.1%),

although the sample size of this species (35 specimens) is

relatively small. Other, mostly non-drilling predatory gas-

tropods that are common in Groß Pampau, and are

known to have molluscs among their prey in modern

communities (Table 2and references therein), belong to

the neogastropod clades Buccinidae (Colus), Conidea

(Conilithes), Fasciolariidae (e.g. Aquilofusus and Latirulus)

and to Retusidae (Pyrunculus). However, owing to the

lack of distinct signatures of predation in the prey’s shells,

it is not feasible to estimate the predation intensity

exerted by non-drilling predatory gastropods from the

fossil record. We tentatively infer that it is quite likely

that the continuous and frequent presence of predatory

gastropods (Fig. 11B) exerted appreciable predation pres-

sure on the Miocene molluscan assemblages of Groß

Pampau, albeit at a level that cannot be assessed more

precisely. Owing to the stabilizing role of predators, we

hypothesize that predatory gastropods exerted a top-down

control on the communities, contributing to the relatively

high taxonomic diversity and stability of the Groß Pam-

pau faunal associations.

An alternative or additional mechanism that would

explain the stable species composition is ecological incum-

bency. Ecological incumbency or priority effects consist of

competitive interspecies-level interactions, in which the

incumbent taxa are advantaged over newly entering taxa

because their population is numerically stronger than that

of the invading species (Jablonski & Sepkoski 1996). This

is especially the case when priority effects derive from

niche pre-emption (i.e. the reduction of resources available

to other potentially invading species by early arriving spe-

cies; Fukami 2015). Given that predators can weaken the

competitive ability of prey, high predation pressure is usu-

ally related to decreased priority effects (Chase et al.2009;

Fukami 2015). However, priority effects can also prevent

invasion by niche modification. In niche modification,

incumbent species directly or indirectly modify the type of

niches available in a local site with negative effects on the

ecological conditions required by the newly arriving species

(Fukami 2015). For the Late Miocene benthic ecosystem at

Groß Pampau, a physical modification of the environment

by the ubiquitous burrowing deposit feeders might have

led to ecological incumbency effects on suspension feeders

(trophic group amensalism), thereby favouring the tempo-

ral persistence of faunal associations.

The stability of faunal associations over longer time-

spans is from time to time interrupted by changes to a

somewhat different taxonomic and functional composi-

tion (Fig. 9). These occasional shifts to differently com-

posed faunal associations and, for example, the

conspicuous stratigraphic distribution of the two species

of Gemmula and the two species of Astarte (Fig. 8), can-

not be explained by any obvious change in the abiotic

environment, be it grain size or clay mineral composition

of the substrate or in the geochemical proxy data (Figs 13, 14).

This also holds for the most pronounced shift in species

composition from faunal association A to those of clus-

ter B in assemblage S21 (Fig. 9). Only the inferred eco-

logical adaptations of the bivalves Varicorbula gibba and

Astarte vetula, both common elements of A but rare

thereafter, hint at a somewhat less favourable environ-

ment, possibly with transient hypoxic conditions or ele-

vated turbidity (see above). Thus, the higher proportions

CAROBENE ET AL.: PALAEOENVIRONMENT OF GROß PAMPAU 25

of infaunal suspension feeders in MOL cluster 2 than in

cluster 1 would only be an epiphenomenon, brought

about by two stress-resistant bivalve species that happen

to be suspension feeders. For the other shifts in taxo-

nomic and ecological composition we can only speculate

on whether the observed pattern is related to subtle

changes in environmental conditions, such as the low

variation in TOC content. Alternatively, deterministic

ecological processes (e.g. driven by variations in the

abundance of predatory gastropods; Fig. 11B) may have

been important (Chase et al.2009), although no statisti-

cally significant relationship is evident. Finally, shifts to a

different faunal association might derive from stochastic

biological processes such as vagaries in the recruitment

and dispersal of larvae.

Comparison of Late Miocene molluscan assemblages of the

southern North Sea Basin

Given the similar low-energy offshore settings of Groß

Pampau (northern Germany) and Gram (southwestern

Denmark) and marine connectivity between both sites, it

seems surprising that cluster analysis at the genus level

showed total separation of their molluscan assemblages

(Fig. 12A). Many molluscan genera, mostly gastropods,

were found only at Groß Pampau (23 genera) or at Gram

(14 genera) (Carobene et al.2023, datasheet 3). Although

these 37 genera tend to be rare, differences are also evi-

dent in the relative abundance of genera common to both

localities. Among the abundant genera, Yoldiella is ubiq-

uitous, whereas the percentages of a few prominent

bivalve genera (Limopsis,Cyclocardia,Goodallia,Varicor-

bula and Alveinus) differ distinctly within and between

the two localities.

While Limopsis and/or Cyclocardia are important faunal

elements of Gram associations 1–3, composed entirely of

samples from Gram, they only rarely occur in the assem-

blages of Groß Pampau (0.5% of the total number of

individuals). However, literature and field observations

confirm the presence of Limopsis and Cyclocardia in the

metacommunity of Groß Pampau (Hinsch 1990; Spiegler

€

urs 1996). In the Middle–Upper Miocene mica-clay

deposits of Schleswig-Holstein and Mecklenburg-

Vorpommern, Limopsis and Cyclocardia assemblages are

usually associated with the deepest water conditions

(Hinsch 1987,2000,2001). Therefore, we infer that, at

the time of deposition of the sampled part of the Groß

Pampau succession, the bathymetric position at Gram

was probably slightly deeper than in Groß Pampau. Fur-

thermore, the decrease of Limopsis and Cyclocardia in

Gram associations 4–6, and the concurrent increase of

comparatively shallower water taxa such as Yoldiella,

Goodallia and Astarte, are consistent with a shallowing-

upwards trend in the Gram Formation (Rasmussen 1966;

Piasecki 2005).

The peak abundances of Varicorbula and Alveinus in

Groß Pampau associations 1 and 3 are in contrast to their

virtual absence in Gram BH I. Owing to the opportunistic

character of Varicorbula gibba (Yonge 1946;Hrs-

Brenko 2006), this difference suggests that those assem-

blages in the lower part of the Groß Pampau section that

yielded abundant Varicorbula experienced some degree of

oxygen depletion, whereas oxygenated conditions prevailed

throughout at Gram. For the early Gram clay deposits at



ade (Jutland, Denmark) belonging to the A. vetula Zone,

Rasmussen (1966) inferred unfavourable reducing environ-

mental conditions on the basis of the high pyrite content

and the small-sized shells of molluscan assemblages. How-

ever, no similar qualitative observations were reported

from the molluscan zones of Gram BH I. If the low values

of raw richness at the base of Gram BH I are due to unsta-

ble conditions, their subsequent increase suggests that

more favourable ecological conditions were eventually

established at Gram as proposed by Piasecki (2005).

Regarding ecologically defined associations (Fig. 12B), all

assemblages from Gram occur in MOL associations that

also contain assemblages from Groß Pampau. This ecologi-

cal congruence shows that despite differences in genus

composition and abundances, the benthic molluscan eco-

systems at both sites operated in a very similar manner.

CONCLUSION

Our multidisciplinary approach, integrating quantitative

sedimentological, geochemical and faunal analyses, pro-

vides an in-depth reconstruction of the ‘Glimmerton’

palaeoenvironment at Groß Pampau and the biotic and

abiotic factors controlling its diverse molluscan assem-

blages. The studied 6-m-thick upper part of the fully

marine succession is early Tortonian in age. Small grain

size and clay mineral composition indicate an offshore

depositional setting. The monotonous silty facies and fau-

nal evidence suggest a low-energy habitat, mostly below

storm wave base, at a water depth of several tens of

metres. Constant and relatively high rates of primary pro-

duction are supported by nitrogen isotope values, the off-

set in d

C composition between benthic and planktonic

species and the high proportion of deposit feeders in the

macrobenthic assemblages. Along with land-derived

organic matter, high primary productivity apparently

maintained a complex marine food web. The d

O com-

position of planktonic gastropod shells indicates a mean

surface water temperature of 18.7°C and 19.8°C depend-

ing on the value used for standard mean ocean water,

which is in accordance with the warm temperate climate

conditions estimated for the Middle to early Late

26 PAPERS IN PALAEONTOLOGY

Miocene of northern Europe. By contrast, bottom water

temperatures determined from the d

O composition of

benthic gastropods were c.9°C lower on average, result-

ing in a pronounced thermal gradient from surface to

bottom waters and a marked summer thermocline.

Low variability in sedimentological and geochemical

proxies coupled with the relatively uniform taxonomic and

functional composition of the fauna indicate generally sta-

ble and favourable ecological conditions for benthic mol-

luscan communities. Although correlation does not

necessarily indicate causation, statistical analyses suggest

that the most important abiotic environmental factors

influencing the distribution of faunal assemblages seem to

be total organic carbon content and total nitrogen content.

In terms of individual responses of species to environmen-

tal factors, the positive correlation between the opportunis-

tic bivalve species Varicorbula gibba and total nitrogen

concentration suggests occasionally less favourable bottom

conditions, possibly related to transient hypoxia.

The temporal distribution of faunal associations defines

a pattern of punctuated continuity, with taxonomic and

functional composition persisting over time before shift-

ing to a somewhat different configuration, which in turn

remains stable for some time before shifting again. Pre-

vailing stable environmental conditions most likely were

not the only prerequisite for taxonomic and ecological

consistency. We envisage top-down control by numerous

predatory gastropods and/or niche modification by

incumbent deposit feeders as an important additional fac-

tor for faunal stability, whereas the causes of faunal shifts

remain speculative.

Comparison with the Late Miocene molluscan assem-

blages of the palaeoenvironmentally similar succession at

Gram, Denmark, indicates differences in the composition

and relative abundance of constituent genera. However,

the pronounced ecological congruence of their molluscan

assemblages indicates that the offshore macrobenthic eco-

systems of the southern North Sea Basin functioned very

similarly.

Acknowledgements. We are grateful to Ronald Janssen (Sencken-

berg Frankfurt) and Gerhard Stein (L€

ubeck) for taxonomic

advice, the Geological Survey of Schleswig-Holstein for providing

documentation on the Groß Pampau boreholes, and Max Vodel

(Museum f€

ur Natur und Umwelt) for support in the field and

processing of samples. We are indebted to Wolfgang Ohle and

Bernard Lau for access to the pit; and to the excavation team

members, Gerhard H€

opfner, Wolfgang H€

opfner, Uwe Havekost,

Andreas Malchow, Martin Kupsch and Svenja Warnke, for sup-

port during field work. Many thanks to Ole Strehlow (Berlin) for

performing part of the sedimentological analyses, Elke Siebert

(Museum f€

ur Naturkunde Berlin) for improving figure design,

and the MfN-Digitization-Team with Michael Neumann

(Museum f€

ur Naturkunde Berlin) for photographic work. We also

thank Michael Hautmann and Sally Thomas for handling the

manuscript and two anonymous journal reviewers for their valu-

able comments and advice. This work was financially supported

by the Gemeinn€

utzige Sparkassenstiftung L€

ubeck and the Centre

of Cultural Research L€

ubeck (University of L€

ubeck). This is

Paleobiology Database official publication number 455. Open

Access funding enabled and organized by Projekt DEAL.

Author contributions. Conceptualization D Carobene (DC), M

Aberhan (MA); Data Acquisition DC, R Bussert (RB); Data

curation DC, MA, CJ Reddin (CJR); Formal Analysis DC, U

Struck (US), RB; Investigation DC, MA, US, RB, CJR; Method-

ology DC, MA, CJR; Project Administration DC, MA;

Resources DC, MA; Supervision MA; Visualization DC;

Writing –Original Draft Preparation DC, MA, US, RB, CJR;

Writing –Review & Editing DC, MA, RB, CJR.

DATA ARCHIVING STATEMENT

Data for this study are available in the Dryad Digital

Repository: https://doi.org/10.5061/dryad.0p2ngf253.

Editor. Michael Hautmann

SUPPORTING INFORMATION

Additional Supporting Information can be found online (https://

doi.org/10.1002/spp2.1496):

Appendix S1. Data sources for Table 2.

Fig. S1. Heat-map diagram of a two-way hierarchical cluster

analysis of Late Miocene faunal assemblages (Q-mode) and 112

benthic macroinvertebrate species (gastropods, bivalves, scapho-

pods, and brachiopods) (R-mode) from Groß Pampau based on

the relative abundances of species. Statistically distinct clusters of

assemblages S1–S30 define six faunal associations. Colour coding

indicates the relative abundance of species.

Table S1. Late Miocene macrobenthic associations of Groß

Pampau based on cluster analysis (Fig. 5) with relative abun-

dance and presence percentage of species attaining at least 1%

relative abundance.

Table S2. Late Miocene functional associations of Groß Pampau

based on cluster analysis (Fig. 7) with relative abundance and

presence percentage of modes of life.

Table S3. Late Miocene molluscan associations of Gram bore-

hole I zones I–IV (GB) and Groß Pampau (GP) based on cluster

analysis (Fig. 12A) with relative abundance and presence per-

centage of genera attaining at least 1% relative abundance.

Table S4. Late Miocene functional associations of Gram bore-

hole I zones I–IV (GB) and Groß Pampau (GP) based on cluster

analysis (Fig. 12B) with relative abundance and presence per-

centage of modes of life.

Table S5. The top six rows show the biplot scores of the envi-

ronmental variables against the CCA dimensions, followed by

eigenvalues and cumulative proportions explained by the CCA

CAROBENE ET AL.: PALAEOENVIRONMENT OF GROß PAMPAU 27

dimensions and their significance as tested by the ANOVA-like

permutation test.

Table S6. Spearman’s rho and significance level of the correla-

tion between the most abundant species and environmental

variables.

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