No Job Name


Foraminiferal morphogroups in dysoxic shelf deposits from
the Jurassic of Spitsbergen
Jenö Nagy,1 Matías Reolid2 & Francisco J. Rodríguez-Tovar3

1 Department of Geosciences, University of Oslo, P.O. Box 1047, Blindern, NO-0316 Oslo, Norway

2 Departamento de Geología, Campus Las Lagunillas, Universidad de Jaén, ES-23071 Jaén, Spain

3 Departamento de Estratigrafía y Paleontología, Universidad de Granada, Avenida Fuentenueva, ES-18071 Granada, Spainpor_112 214..221

Abstract

Analysis of benthic foraminiferal assemblages was performed in Bathonian to
Kimmeridgian deposits through a section covering the lower half of the
Agardhfjellet Formation in central Spitsbergen. The section consists mainly of
organic-rich shales, which contain low-diversity agglutinated assemblages. In
this foraminiferal succession five morphogroups were differentiated according
to shell architecture (general shape, mode of coiling and number of cham-
bers), integrated with the supposed microhabitat (epifaunal, shallow infaunal
and deep infaunal) and feeding strategy (suspension-feeder, herbivore,
bacterivore, etc.). The environmental evolution of the analysed section is
interpreted by using the stratigraphic distribution of morphogroups, com-
bined with species diversities and sedimentary data, in a sequence
stratigraphic framework. The section comprises two depositional sequences,
which demonstrate that species diversity and relative frequency of morpho-
groups are correlative with transgressive–regressive trends controlling depth
and oxygenation of the water column. In both sequences, the maximum
flooding interval is characterized by increased organic carbon content, domi-
nance of the epifaunal morphogroups and reduced species diversity: features
reflecting the increased degree of stagnation separating the transgressive
phase from the regressive phase.

Keywords
Foraminifera; Jurassic; morphogroups;

microhabitats; palaeoenvironments;

Spitsbergen.

Correspondence
Jenö Nagy, Department of Geosciences,

University of Oslo, P.O. Box 1047, Blindern,

NO-0316 Oslo, Norway. E-mail:

jeno.nagy@geo.uio.no

doi:10.1111/j.1751-8369.2009.00112.x

As an approach to palaeoenvironmental assessment,
benthic foraminiferal morphogroup analysis has attracted
special attention over the last two decades. Following
initial papers focused on the palaeoecological significance
of foraminiferal morphology (Chamney 1976; Severin
1983), the morphogroup concept was introduced by
Jones & Charnock (1985). The concept was further devel-
oped in subsequent studies, and has been applied to
interpret various aspects of the palaeoenvironment, such
as food supply, oxygenation, substrate and salinity (Nagy
1992; Tyszka 1994; Bąk 2004; Szydlo 2004; Reolid et al.
2008; among others). The morphogroup approach in this
paper is applied to a foraminiferal succession formed
in mainly dysoxic environments, in order to recognize
changing depositional conditions in a sequence strati-
graphic framework.

The analysed deposits

Jurassic shelf environments characterized by changing
oxygen levels are well represented in the Agardhfjellet
Formation, which is extensively exposed in the Central
Basin of Spitsbergen (Dypvik et al. 1991). This study deals
with the lower 108 m of the Agardhfjellet Formation,
comprising Bathonian, Callovian, Oxfordian and Kim-
meridgian deposits, which have been logged and sampled
from the Janusfjellet section, located in central Spitsber-
gen (Figs. 1, 2).

The lower part of the analysed interval corresponds
to the Oppdalen Member, which begins with the
Brentskardhaugen Bed, a transgressive conglomerate
containing phosphatic pebbles. This is overlain by the
Marhøgda Bed, consisting of ooids in a sandy matrix. The

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mailto:nagy@geo.uio.no


succeeding main body of the member reveals a fining-
upward development from fine sand and silt to shale. The
Lardyfjellet Member consists mainly of finely laminated
black shales with extremely high organic carbon content.
These beds are overlain by the Oppdalssåta Member,
which consists of dark silty shales interbedded with thin
horizons of fine-grained, highly bioturbated sandstones.
The upper part of the analysed section corresponds to the
lower half of the Slottsmøya Member, consisting of lami-
nated dark shales and grey silty shales.

Materials and methods

The succession studied preserves an essentially
continuous benthic foraminiferal record, from which
approximately 6700 specimens have been counted, in
samples taken from 31 levels. Most of the samples were
collected from well-indurated black shales that charac-
terize the main body of the Agardhfjellet Formation. A
smaller number of samples were taken from siltstones
and fine-grained sandstones from the basal part of the
formation. In the laboratory, the rock samples were dis-
integrated using the kerosene method described by
Nagy et al. (1988). The most critical phase of the pro-
cedure is the sediment disintegration, which involves
soaking the sediment in kerosene, followed by boiling
the partly disintegrated sample in sodium hydroxide. A
large number of shale samples required between one
and three repetitions of this step of the procedure (com-
bined with hydrogen peroxide) to obtain an adequate
degree of disintegration.

Morphogroups and subgroups

A detailed analysis of the studied succession revealed a
high abundance of agglutinated foraminifera, and the

almost total absence of calcareous benthic specimens. In
total, 35 agglutinated and two calcareous species have
been observed. Calcareous forms occur in very few
samples, where they compose <1% of the assemblage.
In the foraminiferal succession, five morphogroups are
distinguished and are designated alphabetically from A
to E (Fig. 3; Appendix). Two of these groups, C and D,
are further subdivided into three and two subgroups,
respectively. The differentiation of morphological cat-
egories is based on three types of criteria: (1) test
morphology, comprising general outline, number of
chambers, chamber arrangement and architecture; (2)
life habitat, characterized as deep infaunal, shallow
infaunal or epifaunal; and (3) supposed feeding
strategies, characterized as suspension feeders, deposit
feeders, herbivores or bacterial scavengers. A useful
basis for the assignment of Jurassic genera to morpho-
logical categories and their environmental interpretation
has accumulated as a result of several studies: Nagy
(1992), Tyszka (1994), Nagy et al. (1995) and Reolid
et al. (2008).

A morphogroup system developed for a given sedi-
mentary succession is often cumbersome to apply
directly to a new sedimentary succession, owing to
major differences in the composition of assemblages,
which are mainly determined by divergences in ecosedi-
mentary conditions, and, commonly, differencences in
stratigraphic age. This is also the case with the aggluti-
nated foraminiferal succession of the Jurassic of
Spitsbergen, which required the development of a
modified morphogroup system suitable for the analysis
of assemblages adapted to hypoxic marine shelf envi-
ronments. The typical species of the Svalbard
morphogroups are illustrated in Fig. 4. In addition, illus-
trations of common species occurring in the section
analysed are illustrated in Nagy & Basov (1998).

Fig. 1 Location map of Spitsbergen, Svalbard,
showing the position of the studied section at

Janusfjellet.

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Morphogroup A

This morphogroup comprises species with tubular and
unilocular shells. An epifaunal habitat and suspensivo-
rous feeding strategies are supposed. The group is
represented only by Rhizammina (Fig. 4a), and is equiva-
lent to subgroup 1-a of Nagy (1992) and morphogroup
A-1 of Tyszka (1994).

Morphogroup B

This category is represented by unilocular shells of spheric
and subspheric shape. The organisms are interpreted as
epifaunal, passive deposit feeders (Fig. 3), among which
Thuramminoides is the representative genus (Fig. 4b). The
morphogroup is equivalent to subgroup 2-a of Nagy
(1992).

Morphogroup C

This morphogroup comprises species with elongated,
subcylindric, multilocular shells, adapted to infaunal life
habitats (Fig. 3). Three subgroups have been differenti-
ated within the group.

Subgroup C1 consists of uniserial forms living as
shallow to deep infaunal detritivorous bacterial scaven-
gers. Reopax is the representative genus of this subgroup
(Fig. 4c), which is equivalent to subgroup 3-b of Nagy
(1992) and morphogroup A-8 of Tyszka (1994).

Subgroup C2 is composed of species having a planispi-
ral or streptospiral initial stage, changing to a uniserial
final stage. A shallow infaunal habitat and a detritivorous
bacterial scavenger feeding strategy are inferred. Two
genera, Ammobaculites (Fig. 4d) and Bulbobaculites, are
registered in this subgroup. The subgroup corresponds to
subgroup 3-b of Nagy (1992) and morphogroup A-6 of
Tyszka (1994).

Subgroup C3 consists of biserial, triserial and elongated
trochospiral forms, adapted to shallow to deep infaunal
habitats, with detritivorous bacterial scavenger feeding
strategies. The genera included here are Eomarsonella,
Dorothia, Verneuilina and Verneuilinoides (Fig. 4e). The sub-
group correlates to subgroup 3-b of Nagy (1992) and
morphogroup A-8 of Tyszka (1994).

Morphogroup D

This morphogroup includes species with spiral multilocu-
lar shells and epifaunal to shallow infaunal habitats,
combined with herbivorous and detritivorous feeding
strategies. Two subgroups are differentiated.

Subgroup D1 includes globular and plano-convex
forms, with trochospiral coiling. Epifaunal herbivorous,
detritivorous and active omnivorous feeding habits are
interpreted. The registered genera are Ammoglobigerina
and Trochammina (Fig. 4f). The subgroup is equivalent to
subgroup 2-b of Nagy (1992) and morphogroup A-4 of
Tyszka (1994).

Subgroup D2 is typified by rounded, planispiral mor-
photypes, adapted to epifaunal and potentially shallow
infaunal habitats, with herbivorous, detritivorous and
active bacterivorous behaviour. It is represented by the

Fig. 2 Outline of the stratigraphy, lithology and environments of the
Bathonian to Kimmeridgian part of the Agardhfjellet Formation in the

Janusfjellet section, central Spitsbergen. (Lithological column redrawn

and modified from Århus 1998.)

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Polar Research 28 2009 214–221 © 2009 The Authors216



Fig. 3 Morphogroups (and subgroups) dif-
ferentiated in the Jurassic foraminiferal suc-

cession of the Janusfjellet section, with the

interpreted life positions indicated.

Fig. 4 Species exemplifying the foraminiferal morphogroups recognized in the Janusfjellet section: (a) Rhizammina sp., morphogroup A; (b) Thurammi-
noides sp., morphogroup B; (c) Reophax sterkii, subgroup C1; (d) Ammobaculites areniferus, subgroup C2; (e) Verneuilinoides aff. graciosus, subgroup C3;

(f) Trochammina praerosacea, subgroup D1; (g) Recurvoides scherkalyensis, subgroup D2; (h) Ammodiscus sp., morphogroup E. Scale bars: 250 mm.

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genera Haplophragmoides, Cribrostomoides and Recurvoides
(Fig. 4g). This category correlates with subgroup 3-a of
Nagy (1992) and morphogroup A-6 of Tyszka (1994).

Morphogroup E

This category includes flattened tests composed of two
chambers (proloculus and second chamber), with
planispiral or irregular coiling. An epifaunal to phytal
lifestyle with herbivorous and detritivorous trophic
strategies are interpreted. The genera Glomospira, Areno-
turrispirillina and Ammodiscus (Fig. 4h) have been
recognized. The morphogroup is equivalent to subgroup
4-a of Nagy (1992) and morphogroup A-3 of Tyszka
(1994).

Morphogroups of calcareous taxa

The extremely rare calcareous faunal component com-
prising Lingulina and Lenticulina represent flattened
multilocular morphotypes, with a potentially deep
infaunal microhabitat, and depositivorous and grazing
omnivorous trophic behaviours. Tyszka (1994) include
these forms in morphogroups C-6 (Lingulina) and C-8
(Lenticulina).

Morphogroup trends in a
transgressive–regressive framework

The base of the studied section is an erosional unconfor-
mity cutting the underlying Knorringfjellet Formation.
This ravinement surface represents a sequence boundary,
which became flooded by the regional Bathonian
transgression. During the advancing transgression, a
principally fining-upward succession was formed, reflect-
ing the deepening upward depositional environment of
the Oppdalen Member (Fig. 2). It starts with the
Brentskardhaugen Bed, a beach conglomerate containing
reworked phosphatic pebbles, followed by the sandy
oolites of the Marhøgda Bed, which is succeeded by a fine
sand to shale succession, forming the rest of the member.

No foraminifera were found in the lower, coarser part
of the section (Fig. 5). Above this level, the number of
species is relatively low, suggesting the effect of a restrict-
ing factor, possibly reduced salinity in surficial waters.
The development of low salinity in the upper part of the
water column can be explained by large freshwater dis-
charge to the basin from the extensive adjacent land
areas. The effect of this factor decreases upwards in the
Oppdalen Member, as shown by the increasing species
diversity. In the upper part of the member, there is a
diversity reduction owing to decreasing oxygen levels, as
evidenced by the increasing quantities of organic carbon.

The oxygen-depleted conditions in deeper water could be
attributed to the density-stratified nature of the water
column. The foraminiferal assemblages are dominated by
the digging infaunal component (morphogroup C), fol-
lowed by the epifaunal group (mainly B and D1). The
deep-digging group is associated with low organic carbon
content, which is mainly found in the middle part of
the member.

The fining-upward succession of the Oppdalen
Member terminates in the organic-rich black shale inter-
val of the Lardyfjellet Member. These shales are finely
laminated, papery in consistency and show high values of
organic carbon (5.0–11.2%). The foraminiferal assem-
blages reveal minimum values of diversity and a high
dominance of the epifaunal morphogroups B and D1. The
shallow infaunal component (groups C2 and D2) is
strongly reduced, suggesting that the redox boundary was
located close to the sediment–water interface, and that
there was an absence of oxygen in the typical infaunal
microhabitats. These features indicate dysaerobic to
anaerobic bottom water conditions, which are most
pronounced in this interval within the Agardhfjellet For-
mation. The high degree of stagnation is apparently
related to the maximum flooding interval (Fig. 5).

The Oppdalssåta Member of dark silty shales and
interbedded fine-grained sandstones shows a generally
coarsening-upward trend, reflecting the shallowing of the
depositional area during the regressive phase of the first
subordinate sequence of the Agardhfjellet Formation. The
Oppdalssåta Member sandstones were deposited as off-
shore bars (Dypvik et al. 1991), and their extremely high
degree of bioturbation is in accordance with deposition
under oxic conditions. The shales between the sandbars
were deposited under dysaerobic conditions, however, as
shown by their foraminiferal assemblages that are domi-
nated by the shallow infaunal component, and reveal
increased but still low species diversities (Fig. 5). The
expansion of these two faunal parameters and the
reduced organic carbon content indicate increased oxy-
genation, compared with the underlying maximum
flooding interval.

There is a sharp lithological contact between the sandy
Oppdalssåta Member and the dark shales of the overlying
Slottsmøya Member. The contact represents a trans-
gressive surface, introducing the second subordinate
sequence, which constitutes the upper part of the Agar-
dhfjellet Formation (Fig. 5). The transgressive phase of
this sequence is characterized by moderate diversities and
the high dominance of the shallow infaunal and poten-
tially epifaunal morphogroups (C2 and D2). Higher up in
the Slottsmøya Member, a maximum flooding interval is
developed, as shown by the slightly increased stagnation,
indicated by reduced species diversity, an expanded

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Polar Research 28 2009 214–221 © 2009 The Authors218



epifaunal component (mainly morphogroup D1, and sec-
ondarily B) and increased organic carbon content. The
rather weakly developed base of the regressive phase is
signalled by increased diversity and reduced organic
carbon content. This phase continues to the upper
sequence boundary marking the top of the Agardhfjellet
Formation.

Conclusion

Callovian to Kimmeridgian deposits, composing the
lower half of the Agardhfjellet Formation in the Janus-
fjellet section, central Spitsbergen, consist mainly of
shales with high to intermediate organic carbon con-
tent. Deposition of the shales took place mainly under

Fig. 5 The Janusfjellet section, with sedimentary features, morphogroup categories, diversities and oxygenation in a sequence stratigraphic framework.
(Lithological column redrawn and modified from Århus 1998.)

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Polar Research 28 2009 214–221 © 2009 The Authors 219



dysaerobic conditions. This sedimentary succession con-
tains an essentially continuous foraminiferal record,
composed almost exclusively of agglutinated taxa
forming low-diversity assemblages of restricted nature.
The main restricting factor was reduced access to oxygen
in benthic environments, owing to the salinity-induced
stratification of the water column.

In the foraminiferal succession, five morphogroups
(including five subgroups) have been differentiated,
based on morphological features combined with inferred
microhabitat preferences and feeding strategies. Morpho-
group A comprises forms with tubular unilocular shells,
and an inferred epifaunal suspension-feeding habit. Mor-
phogroup B represents globular unilocular forms with an
epifaunal, passive depositivorous mode of life. Morpho-
group C contains three subgroups composed of elongated,
subcylindrical and multilocular taxa, with an infaunal
habitat and detritivorous bacterial scavenging: subgroup
C1 of uniserial forms living in shallow to deep infaunal
microhabitats; subgroup C2 with spiral initial stage and
uniserial final part, living as shallow infauna; and sub-
group C3 of biserial, triserial or elongated trochospiral
forms, with shallow to deep infaunal strategies. Morpho-
group D comprises spiral multilocular species living as
detritivorous epifauna (potentially shallow infauna),
divided into two subgroups: subgroup D1, with globular
and planoconvex shape, and trochospiral coiling, includ-
ing herbivores, detritivores and active omnivores; and
subgroup D2, including rounded planispiral species
with herbivorous, detritivorous and active bacterivorous
behaviour. Morphogroup E comprises flattened taxa,
with a proloculus and a spiral or irregular second
chamber, adapted to epifaunal to phytal, herbivorous and
detritivorous modes of life.

The stratigraphic distribution of the morphogroups,
combined with sedimentary data and species diversities,
have been employed to delineate the palaeoenvironmen-
tal evolution of the Janusfjellet section, related to sea-
level changes in a sequence stratigraphic framework.
Sea-level changes controlled the oxygenation at the dif-
ferent microhabitat depths at, or below, the seabed
surface, as reflected by the variation in morphogroup
proportions and diversity.

In central Spitsbergen, the Agardhfjellet Formation
(Bathonian to Ryazanian in age) comprises a major
depositional sequence, which can be subdivided into two
subordinate sequences. These two sequences are repre-
sented in the Bathonian to Kimmeridgian interval studied
of the Janusfjellet section, although most of the regressive
phase of the upper sequence is not included in this study.
In both sequences the maximum flooding interval is char-
acterized by the increased organic carbon content, the
dominance of epifaunal foraminifera and the reduced

species diversity: these features all indicate increased
stagnation in benthic environments. The transgressive
and regressive phases reveal a reduced organic carbon
content, the dominance of shallow-digging infauna and
an increased diversity.

Acknowledgements

The participation of J. Nagy in this research has been
supported by the Statoil/Hydro VISTA programme. The
contribution of M. Reolid and F.J. Rodríguez-Tovar has
been supported by the projects CGL2005-06636-C0201
and CGL2005-0316/BTE, the EMMI group (RNM-178,
Junta de Andalucía) and the Acción Integrada
30.AI.PO.1300 (University of Granada–University of
Oslo). A grant of the Universidad de Jaén financed
M. Reolid’s short stay at the University of Oslo.

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Appendix: foraminiferal species by
morphogroups and subgroups

Morphogroup A

Rhizammina sp.

Morphogroup B

Thuramminoides lapilliformis Nagy & Basov, 1998; Thuram-
minoides sp.

Morphogroup C

Subgroup C1
Reophax sterkii Haeusler, 1890

Subgroup C2
Ammobaculites areniferus Nagy & Basov, 1998; Ammobacu-
lites suprajurassicum (Schwager, 1865); Ammobaculites sp.;
Bulbobaculites aff. elongatulus (Dain, 1972).

Subgroup C3
Dorothia sp.; Eomarsonella paraconica Levina, 1972; Ver-
neuilina sp.; Verneuilinoides aff. graciosus Kosyreva, 1972;
Verneuilinoides sp.

Morphogroup D

Subgroup D1
Ammoglobigerina canningensis (Tappan, 1955); Trocham-
mina kosyrevae Levina, 1972; Trochammina aff. minutissima
Dain, 1972; Trochammina misinovi Levina, 1972; Trocham-
mina praerosacea Nagy & Basov, 1998; Trochammina
rostovzevi Levina, 1972; Trochammina sp.; Trochammina
spp.

Subgroup D2
Cribrostomoides subretusus Nagy & Basov, 1998; Cribrosto-
moides sp.; Haplophragmoides canuiformis Dain, 1972;
Haplophragmoides sp.; Recurvoides disputabilis Dain, 1972;
Recurvoides aff. disputabilis Dain, 1972; Recurvoides scherka-
lyensis Levina, 1962; Recurvoides aff. scherkalyensis Levina,
1962; Recurvoides sp.

Morphogroup E

Ammodiscus aff. pseudoinfimus Gerke & Sossipatrova, 1961;
Ammodiscus sp.; Arenoturrispirillina jeletzkyi Chamney,
1971; Glomospira oxfordiana Sharovskaja, 1966; Glomospira
semiaffixa Sharovkaja, 1966.

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