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Characteristic trace fossils from Miocene brackish-water deposits in the Niger

Delta, Nigeria

Article  in  Geologos · August 2018

DOI: 10.2478/logos-2018-0011

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 Sunny C. Ezeh, Wilfred A. Mode, Berti M. Ozumba

Geologos 24, 2 (2018): 111–125

doi: 10.2478/logos-2018-0011

Characteristic trace fossils from Miocene brackish-

water deposits in the Niger Delta, Nigeria

Sunny C. Ezeh1*, Wilfred A. Mode1, Berti M. Ozumba2

1
Department of Geology, University of Nigeria, Nsukka, 410001, Enugu State, Nigeria
2
Independent Petroleum Geological Consultant
*corresponding author, e-mail: [email protected]

Abstract

Miocene deposits in the eastern portion of the Greater Ughelli, Central Swamp and Coastal Swamp depobelts contain
well-developed brackish-water trace fossil assemblages. Twelve ichnogenera have been identified, namely: Asterosoma,
Bergaueria, Chondrites, Gyrolithes, Thalassinoides, Lockeia, Palaeophycus, ?Conichnus, Planolites, Siphonichnus, Skolithos and
Diplocraterion. In addition, common non-descript, passively filled burrows and fugichnia (escape structures) have also
been observed. The above-mentioned ichnogenera and associated non-descript structures can be arranged into six dis-
tinct and recurring ichnoassociations within the Greater Ughelli, Central Swamp and Coastal Swamp depobelts. Each
ichnoassociation is comprised of a group of trace fossils which collectively reflect specific environmental conditions
during deposition of these Miocene strata. All trace fossil assemblages illustrate deposition in nearshore, restricted
settings. Ichnological and sedimentological criteria which may be utilized to recognise brackish-water deposits are
discussed and illustrated in pictures of the cores studied.

Key words: Ichnoassociations, ichnology, salinity, Greater Ughelli, Central Swamp, Coastal Swamp, Neogene

1. Introduction
delta’s most prolific play since the onset of explora-
tion activities in the basin over fifty years ago. Ac-
Three geological units in the Niger Delta, i.e., the cording to CSL 2014, the play represents 25 per cent
Akata, Agbada and Benin Formations, came into of all exploration wells (1,300 in number) drilled in
existence during five offlapping siliciclastic sedi- the Niger Delta and 40 per cent of onshore wells
mentation cycles (Ezeh et al., 2016a) otherwise re- (822 in total). The average field size for the onshore
ferred to as depositional belts or depobelts or the Niger Delta is ~ 90 million barrels (MMbbls), with
Northern Delta, Greater Ughelli, Central Swamp, a potential future capacity of sixteen billion barrels
Coastal Swamp and Offshore. Except for the last- (Bbbls) of oil equivalent (10 Bbbls for oil and 6 Bbbls
named, which belongs to the offshore Niger Del- for gas).
ta, all remaining depobelts are located within the The Akata Formation represents the oldest unit,
onshore portion of the Niger Delta Basin. Miocene consisting of marine shales, and is overlain by the
strata have remained essential in research, especial- paralic Agbada Formation which represents the
ly of the onshore Niger Delta, owing to their being main reservoir unit. Most of the oil discovered
the primary constituent the central-eastern portion in the delta is found in this particular unit which
of the Greater Ughelli, Central Swamp and Coastal comprises intercalations of sandstone and shale se-
Swamp depobelts (Fig. 1). It has been amongst the quences. The sandstones constitute an essentially
112 Sunny C. Ezeh, Wilfred A. Mode, Berti M. Ozumba

Fig. 1. Map of the study area, show-

ing the five main depositional
belts (depobelts) and the wells
in the Niger Delta sedimentary
basin studied.

poorly consolidated reservoir, while the shales also The aim of the present paper is to reconstruct
function as source rocks and cap rocks (Michele et the palaeoenvironment of Miocene deposits in the
al., 1999). On top of the Agbada Formation follows Niger Delta, based on trace fossil assemblages of
the Benin Formation which bears little oil and con- the Greater Ughelli, Central Swamp and Coastal
sists mainly of continental sands. Swamp depobelts, which have been useful in inter-
Considering the vast reserves contained within pretations of depositional environments and may
the Greater Ughelli, Central Swamp and Coastal offer valuable input to the development of integrat-
Swamp, this is an important target for further res- ed facies models that could be used to formulate ef-
ervoir studies. However, successful economic ex- fective developmental strategies.
ploitation of these resources requires a thorough
understanding of the sedimentological, stratigraph-
ical and structural controls that have an influence on 2. Geological setting and stratigraphy
hydrocarbon distribution. Consequently, detailed
and integrated facies models are an essential tool The Niger Delta sedimentary basin is situated in
in the most efficient development of the depobelts southern Nigeria, with an areal coverage of 300,000
and ichnological studies have proved to be particu- km2, including the geological extent of the Neogene
larly valuable in palaeoenvironmental interpreta- Niger Delta (Akata-Agbada) Petroleum System
tions and, by extension, in integrated facies models. (Michele et al., 1999). It is bounded in the north and
Unfortunately, there are no previous detailed ich- northeast by the Benin flank, the Anambra Basin
nological studies for the basin, with the exception and the Abakaliki fold belt, in the east-south-east
of recent ichnological work by Ezeh et al. (2016a, by the Calabar flank, while the Cameroon volcanic
b) which characterised shoreface variability, sedi- line lies in the east. The western axis is formed by
mentology and ichnofacies of the Coastal Swamp the Dahomeyan embayment which is the eastern-
and shallow Offshore depobelts. Ezeh et al. (2016a) most West African transform-fault passive bound-
used the fully marine trace fossils recognised to ary (Fig. 2).
characterise and interpret the deposits in the shore- The structure of the continental boundary along
face section, while Ezeh et al. (2016b) showed the the west coast of Equatorial Africa is influenced by
usefulness of trace fossils and cores in the interpre- the Cretaceous fracture zones portrayed as trench-
tation of reservoir facies against the conventional es in the deep Atlantic. These fracture zones ridges
use of well log signatures. Although these recent separate the boundary into different basins and, in
papers have been very useful in our understanding Nigeria, develop the boundary faults of the Creta-
of the depositional history of the area, there still is a ceous Benue-Abakaliki Trough which extend into
need to document the ichnological characteristics as the West African shield. The trough marks the
these relate to brackish-water deposits. failed arm of a rift triple junction linked with the
 Characteristic trace fossils from Miocene brackish-water deposits in the Niger Delta, Nigeria 113

Fig. 2. Geological map of Nigeria with the main sedimentary basins, the basement complex and the prograding Niger
Delta Basin (selected dotted shape) with selected growth faults (adapted after Whiteman, 1982).

opening of the South Atlantic. Lehner & De Ruiter Guinea. They show synsedimentary faulting which
(1977) proposed that the rifting started during the occurred as a result of the interaction of sediment
Late Jurassic and continued into the mid-Creta- supply and subsidence (Doust & Omatsola, 1990).
ceous; however, in the Niger Delta region, rifting The Greater Ughelli, Central Swamp and Coast-
receded entirely in the Late Cretaceous. al Swamp depobelts (latitude 6°62'40"–7°45'00"N;
After rifting ceased, gravity tectonics became the longitude 4°58'12"–5°39'10"E; see Fig. 1) are char-
primary deformational process. Mobile shale gener- acterised by growth faults and associated roll-over
ated internal deformation and occurred in response anticlines. These formed contemporaneously with
to two processes (Kulke, 1995), i.e., shale diapirs and deposition and represent the structural trapping
slope instability. Shale diapirs were formed from mechanism of petroleum in these depobelts.
the loading of poorly compacted, over-pressured, As previously mentioned: the stratigraphy of the
prodelta and delta-slope clays (Akata Formation) Niger Delta, and that of the study area in particular,
by higher-density delta-front sandstone (Agbada can be divided into three major units that range in
Formation), while slope instability developed as a age from Eocene to Holocene (Fig. 3). The Akata
result of a lack of lateral, basinward, support for Formation comprises at least 6,500 metres of marine
the under-compacted delta slope Agbada Forma- clays with silty and sandy interbeds (Whiteman,
tion (Ezeh et al., 2016a). It is believed that gravity 1982), while the Agbada Formation is characterised
tectonics ended prior to the development of the by paralic to marine coastal and fluvial-marine de-
continental Benin Formation; this is characterised posits, composed mainly of sandstones and shales
by complex structures such as shale diapirs, roll- illustrating coarsening-up offlap cycles (Weber,
over anticlines, collapse growth fault crests, back- 1987). The Benin Formation consists of continental
to-back features and steeply dipping, spaced frank and fluvial sands, gravel and back swamp deposits,
faults (Evamy et al., 1978; Xiao & Suppe, 1992). as is 2,500 m in thickness (Reijers, 2011). These three
As stated by Stacher (1995), the depobelts are 30– diachronous formations occur within growth-fault-
60 kilometres in width and prograde 250 kilometres bounded sedimentary units referred to as depobelts
southwestwards over oceanic crust into the Gulf of or depocentres that succeed each other in a south-
114 Sunny C. Ezeh, Wilfred A. Mode, Berti M. Ozumba

Fig. 3. Stratigraphy of the Neogene Niger Delta

sedimentary basin with respective ages of the
formations, facies and wells studied (adapted
after Doust & Omatsola, 1990).

erly direction (Doust & Omatsola, 1990; Stacher, Coastal Swamp depobelts during a period of accu-
1995). The sedimentation in each depobelt (Fig. 1) mulation of littoral and lower coastal-plain deposits
is a function of the rates of deposition and of sub- in the northern delta. In the same vein, Oyanyan et
sidence with syn-sedimentary growth faults upset- al. (2013) identified ten lithofacies from core samples
ting the balance (Evamy et al., 1978). The growth in the course of determining the depositional envi-
faults are generated by rapid sedimentation and ronment of the eastern part of the Greater Ughelli.
gravitational instability during the accumulation of These samples aided in the reconstruction of five
the Agbada deposits and continental Benin sands sub-depositional environments (i.e., lower shore-
over mobile and under-compacted Akata prodel- face, middle shoreface, distributary channel, tidal
ta shales. Lateral flowage and extrusion were also flat and tidal channel) in the Greater Ughelli depo-
responsible for the diapiric structures on the conti- belt. Similarly, Okengwu & Amajor (2014) recog-
nental slope of the Niger Delta (Reijers et al., 1997). nised successions of well-developed shoreface de-
posits, illustrating the combined influence of wave
and fluvial processes and sediments consisting of
3. Previous research in the study area prograding wave-dominated shoreface with a fluvi-
al predominance in the Greater Ughelli and Central
Miocene strata assigned to the Agbada Formation Swamp depobelts. Prince & Minapuye (2015) identi-
represent the largest petroleum-bearing unit of the fied three environments (viz., tidally influenced flu-
Greater Ughelli, Central Swamp and Coastal Swamp vial channel, upper shoreface and lower shoreface)
depobelts. In spite of a vast resource potential there in the stratigraphy of the Greater Ughelli depobelts.
are only very few studies of trace fossil character- Of a more recent date are detailed interpretations of
istics and of the depositional environments of res- depositional environments on the basis of cores of
ervoirs by using core data. In the Coastal Swamp onshore and shallow offshore depobelts by Ezeh et
depobelt, Egbu et al. (2009) observed five lithofacies al. (2016a, b).
associations (i.e., foreshore, upper shoreface, mid-
dle shoreface and lower shoreface) using sedimen-
tological features. While studying the stratigraphy 4. Methodology
and sedimentology of the Niger Delta, Reijers (2011)
noted that shoreface sediments had formed simulta- Detailed core analysis (both sedimentologically and
neously in the Greater Ughelli, Central Swamp and ichnologically) was conducted for three wells (Fig.
 Characteristic trace fossils from Miocene brackish-water deposits in the Niger Delta, Nigeria 115

1) in different fields within the Greater Ughelli (well provide clues with respect to the nature of the dep-
1), Central Swamp (well 2) and Coastal Swamp (well ositional environment. The stratigraphical recur-
3) depobelts. Morphologies of both dominant and rence of these ichnoassociations within the Miocene
associated trace fossils aided in the classification of deposits reflects the re-establishment of particular
ichnoassociations. Trace fossils were observed and environmental conditions throughout deposition
assigned to respective ichnofacies types based on of the Greater Ughelli, Central Swamp and Coastal
Seilacher’s model (1967), as modified by Pember- Swamp depobelts. In general, six ichnoassociations
ton et al. (2001). The degree of bioturbation is based characterise the Miocene deposits within these de-
on Droser & Bottjer’s (1986) ichnofabric index, as pobelts and represent a continuum of palaeoenvi-
modified by Pemberton et al. (1992). Altogether, a ronmental conditions in an overall brackish-water
total length of 334 metres of core has been studied. depositional environment. The present paper out-
Descriptions of sedimentary features and of charac- lines ichnological and sedimentological criteria
teristics of associated trace fossils were used to in- that could be integrated alongside other data so as
terpret the depositional environments. The map of to obtain a better interpretation of the depositional
the study area was developed using ArcGis Map 10. settings. In addition, it helps in gaining an appreci-
ation of the degree of variability of the latter.

5. Results
5.1. Recurring ichnoassociations
Twelve ichnogenera have been recognised in
the Greater Ughelli, Central Swamp and Coastal As previously noted, the overall distribution of
Swamp depobelts, namely Asterosoma, Thalassi- trace fossils within the Miocene sedimentary rocks
noides, Bergaueria, Gyrolithes, Chondrites, Palaeophy- of the Greater Ughelli, Central Swamp and Coast-
cus, Siphonichnus, Skolithos, Planolites, Lockeia, Dip- al Swamp depobelts can be viewed in terms of six
locraterion and ?Conichnus. In addition to distinct distinct, recurring ichnoassociations (Table 2). The
trace fossils, bioturbated textures, escape structures concept of recurring ichnoassociations has seen ex-
and non-descript, passively filled shafts have also tensive use in palaeoecological and sedimentolog-
been noted. Although slightly problematic, each ical interpretations. The basis for this recurrence
ichnogenus can be attributed to a particular group stems from the concept that trace fossils constitute
(or groups) of organisms, to an ethological (or be- the preserved record of behaviour and functional
havioural) category and, lastly, to a general trophic morphology and, therefore, reflect adaptations of
group (Table 1). organisms to particular ecological conditions (Frey
The twelve ichnogenera recognised, together & Pemberton, 1985; Beynon & Pemberton, 1992).
with associated non-descript structures and tex- Ichnoassociations characteristic of particular envi-
tures, have been seen to occur in six distinct ich- ronmental regimes are recurrent in space and time
noassociations. Each trace fossil association is com- whenever the requisite environmental conditions
prised of a number of ichnogenera. Collectively, the occurred (Frey & Seilacher, 1980; Frey & Pember-
constituent trace fossils of each ichnoassociation ton, 1985).

Table 1. Ethological classification of brackish-water trace fossils from the Greater Ughelli, Central Swamp and Coastal
Swamp depobelts
Trace fossil Ethological classification Trophic strategies Probable organism
Gyrolithes Domichnia deposit-feeder Annelid
Bergaueria Domichnia/Cubichnia suspension feeder Anemone
Palaeophycus Domichnia carnivore Annelid
Siphonichnus Fodinichnia suspension feeder Annelid
Asterosoma Fodinichnia deposit-feeder Annelid/Crustacean
Planolites Fodinichnia/Pascichnia deposit-feeder Annelid
Lockeia Cubichnia deposit-feeder Bivalve
Chondrites Fodinichnia/Chemichnia/Agrichnia deposit-feeder Siphunculid/Annelid
Thalassinoides Fodinichnia/Domichnia deposit-feeder Decapod/Crustacean
Conichnus Domichnia/Cubichnia carnivore/suspension feeder Sea anemone
Skolithos Domichnia suspension feeder Annelid
Diplocraterion Domichnia/Equilibrichnia suspension feeder Crustacean
116 Sunny C. Ezeh, Wilfred A. Mode, Berti M. Ozumba

Table 2. Distribution and relative abundance of ichnogenera in recurrent brackish-water trace fossil assemblages in
Miocene deposits within the cores of the Greater Ughelli, Central Swamp and Coastal Swamp depobelts.
Ichno-association Gyrolithes-Chondrites Planolites Skolithos Palaeophycus Asterosoma Thalassinoides
Trace fossil
Gyrolithes c–a – – – – –
Bergaueria – – – – – o
Palaeophycus – r – o–r c o–r
Siphonichnus – – – c–a – –
Asterosoma – – – – – c
Planolites c c – c c r
Lockeia o – – – – –
Chondrites c–a r – – – –
Thalassinoides r r – – r a
Conichnus – – – – – *o–r
Skolithos – – a – – –
Diplocraterion – – – – o –
Depobelts CS GU & Centr.S GU & CS Cent.S GU CS
c – common, a – abundant, r – rare, o – occasional, * – opportunistic trace fossil, CS – Coastal Swamp Centr.S – Central
Swamp GU – Greater Ughelli.

5.1.1. Gyrolithes-Chondrites ichnoassociation are abundant in the middle portions (4,051–4,054 m

This association is characterised by a low-diversi- depth) of well 3 in the Coastal Swamp depobelt. Bi-
ty ichnoassemblage which is dominated by Gyro- oturbation intensity is generally low, but a unique
lithes and Chondrites (Fig. 4). Associated ichnofossil feature of this ichnoassociation is the occurrence
taxa and structures may include Lockeia, syneresis of horizons which are pervasively bioturbated by
cracks and load structures. For the most part, this these trace fossils. The low-diversity nature of this
ichnoassociation is restricted to inclined heterolith- ichnoassociation may reflect extremely stressful
ic, cross-stratified sediments and mudstones which environmental conditions such as brackish-water

Fig. 4. Gyrolithes-Chondrites ichnoassoci-

ation (well 3; depth 3,950 m). A – Gy-
rolithes (Gy) within a dark brownish
grey mudstone-heterolithic deposit; B
– Gyrolithes (Gy), Chondrites (Ch) and
syneresis cracks (Sy) on an inclined het-
erolithic, cross-stratified (IHCS) bed; C
– Lockeia (Lk), Chondrites (Ch) and load
structure on an IHCS bed (well 3; depth
3,950.5 m). Scale bar in centimetres.
 Characteristic trace fossils from Miocene brackish-water deposits in the Niger Delta, Nigeria 117

and/or other ecological conditions. The morpholo- 5.1.2. Planolites ichnoassociation

gy of Gyrolithes is interpreted to reflect a burrowing This ichnoassociation is mostly restricted to car-
adaptation to escape from extreme salinity fluctu- bonaceous and laminated mudstones, but locally
ations at the sediment-water interface (Gernant, is found within interlaminated mudstones. The
1972; Powell, 1977; Netto et al., 2007). Gernant relative degree of bioturbation is extremely low,
(1972) suggested that the Gyrolithes trace-making being represented by a low-density, monospecif-
organism was restricted to marginal marine stra- ic assemblage of Planolites (Fig. 5). Accessory fea-
ta and therefore Gyrolithes might be utilized as a tures include abundant carbonaceous debris, both
brackish-water indicator. Although palaeoenvi- as laminae and as disseminated matter as well as
ronmental interpretations constructed on the basis siderite concretions. The relatively low intensity
of only a single sedimentary structure should be of bioturbation suggests that bottom waters were
avoided, it is interesting to note that other lines of not sufficiently oxygenated to support a diverse,
sedimentological evidence such as syneresis cracks oxygen-dependent benthic community. Oxygena-
and current beddings (Fig. 4) are indicative of a tion of the water column is important to all benthic
brackish-water depositional environment, suggest- organisms; however, some taxa can tolerate and
ing this assumption may have some validity in this may even preferentially inhabit low oxygen or ox-
particular case. ygen-depleted ecological niches (Ekdale, 1988). The

Fig. 5. Planolites ichnoassociation. A – Section through an estuarine dark brown-grey laminated and carbonaceous
mudstone with abundant concretions (Cn) and Planolites (well 1; depth 4,031.2–4,034.9 m); B – Laminated mudstone
with concretions (Cn). Note the monospecific nature of trace fossils and their low density. Scale bar in centimetres;
each core box is 0.91 m in length.
118 Sunny C. Ezeh, Wilfred A. Mode, Berti M. Ozumba

inferred nature of the Planolites trace-making organ- tions. The formation of siderite is enhanced when in-
ism (i.e., endostratal deposit-feeding) suggests that, terstitial pore waters are depleted with respect to free
although sediments may have been organic rich, oxygen and dissolved sulphur. Thus, the presence of
the interstitial environment was not completely de- siderite may be suggestive of rapid accumulation
void of oxygen. Thinly laminated zones devoid of and decomposition of organic matter in a restricted,
biogenic structures imply that periodically, anoxic anoxic or oxygen-limited environment (Gauthier,
conditions may have been established. Prolonged 1982). However, the low diversity and density of bi-
periods during which the interstitial environment ogenic structures, abundance of siderite and the car-
and/or the overlying water column were oxygen bonaceous nature of the sediments suggest that the
depleted would have been lethal to benthic organ- interstitial waters were not well oxygenated.
isms (Ekdale, 1985; Beynon & Pemberton, 1992). In turn, this suggests that both salinity and ox-
The distinct lack of biogenic structures created ygenation may have been limiting factors in the
by suspension-feeding organisms implies that the distribution of benthic organisms. It is difficult to
physical and ecological conditions were not con- discern whether or not these factors independent-
ducive to the development of a suspension-feeding ly or dependently influenced the distribution of
community. In low-oxygen environments, sedi- benthic organisms. Regardless of which was the
ments commonly contain high concentrations of un- dominant factor, the low-density, monospecific as-
oxidised organic matter which may support dense semblages indicate that only a limited number of
deposit-feeding communities. Such environments benthic organisms could flourish in this unstable or
are typically stagnant and devoid of currents of suf- unpredictable setting (Beynon & Pemberton, 1992).
ficient strength to suspend nutrients in the water Organisms which inhabit unstable settings, such as
column. As a result, low-oxygen environments are estuaries and other marginal marine environments,
typified by a predominance of deposit-feeding or- naturally have broad environmental tolerances and
ganisms and a general absence of suspension-feed- can adapt to environmental disturbances. Such or-
ing organisms (Tognoli & Netto, 2003). Therefore, ganisms, because they are subjected to high levels of
oxygen-depleted deposits characteristically contain physiological stress, tend to display opportunistic
low-density, low-diversity trace fossil assemblages or r-selected population dynamics (Ekdale, 1985).
that are dominated by deposit-feeding structures. In contrast, benthic organisms which inhabit stable
Alternatively, other environmental factors such as or predictable settings are more severely affected
variable and low salinities may also have imposed by physical or biological stresses such as variable
severe physiological stresses on benthic organisms salinity or oxygenation levels. Conditions of abun-
and have resulted in the preferential exclusion of dant organic matter, fluctuating salinity and low
suspension-feeding organisms. Burrowing, espe- concentrations of dissolved oxygen are common in
cially deep burrowing, is an adaptation which lends estuarine and delta plain environments in which
benthic organisms the capability of withstanding large volumes of organic-rich mud are laid down in
salinity fluctuations due to the buffering capacity restricted settings.
of sediments (Wightman et al., 1987). Sanders et al.
(1965) investigated salinity fluctuations in the water 5.1.3. Skolithos ichnoassociation
and sediments of the Pocasset River Estuary from This association occurs in fine-grained, cross-strat-
2.3 ppm to 29.3 ppm during a single tidal cycle and ified and planar-laminated sandstones, being char-
found that the interstitial environment (depth = acterised by Skolithos in moderate density and low
5–20 cm) remained relatively constant at 20.5 ppm. diversity (Fig. 6). This consists predominantly of
Therefore, even shallowly burrowing organisms vertical dwelling and feeding structures such as
are significantly removed from the harsh physical Skolithos, Siphonichnus and Planolites created by
and chemical environment of the sediment-water suspension-feeding organisms. The predominance
interface and overlying water column. Thus, be- of biogenic structures created by suspension-feed-
cause of the ability of deposit-feeding organisms ing organisms is suggestive of nutrient-rich and
to flourish within the interstitial environment, they well-oxygenated bottom waters. Currents were of
may have been able either to escape from or tolerate sufficient magnitude to suspend nutrients within
harsh ecological conditions and may have preferen- the water column, but were moderate enough to
tially inhabited a stressful ecological niche because allow organic detritus to settle from suspension,
of low levels of interspecific competition (Ekdale et offering nutrient resources to shallow suspen-
al., 1984). sion-feeding organisms. The characteristics of this
The abundance of siderite concretions comple- ichnoassociation, i.e., the predominance of vertical
ments the interpretation of anoxic or reducing condi- dwelling structures, generally low diversity and
 Characteristic trace fossils from Miocene brackish-water deposits in the Niger Delta, Nigeria 119

Fig. 6. Skolithos ichnoassociation. A, B – Skolithos- (Sk) dominated, laminated-cross-stratified sandstone bed (well 3;
depth 2,974 m). Note the moderate density and monospecific characteristics of the ichnoassociation; C, D – Siphon-
ichnus (Si) and Planolites (P) on moderately bioturbated, sandy heteroliths. Similarly, note low diversity, moderate
density and associated mud couplets.

moderate burrow density are, for the most part, in- storm wave base and possibly fairweather wave
dicative of the Skolithos ichnofacies (Beynon & Pem- base. Emplacement of these beds does not represent
berton, 1992; Pemberton et al., 2001). sudden shallowing events, but rather a temporary
Units characterised by this ichnoassociation typ- lowering of wave base in response to coastal up-
ically consist of alternating bioturbated and lami- welling associated with meteorological disturbanc-
nated or cross-stratified beds. The bioturbated beds es. The occurrence of these dwelling structures may
contain a low-diversity, Skolithos-dominated assem- represent the displacement of the resident benthic
blage. Laminated zones reflect energetic periods in community by high physiological stress, associated
which shear stress in the benthic boundary layer with the deposition of sandstone beds (flood event).
could not be tolerated by the benthic community. With the return to normal, fairweather conditions
Bioturbated beds record periods in which current the resident deposit-feeding community was re-es-
velocities were sufficiently reduced, permitting or- tablished and the opportunistic, suspension-feed-
ganisms to rework the substrate. ing community was eventually displaced. Similar
event-related examples of ichnological dynamics
5.1.4. Palaeophycus ichnoassociation have been documented from ancient shallow-ma-
This association is typical of laminated to bioturbat- rine deposits (compare Frey & Seilacher, 1980; Pem-
ed mudstones and heteroliths and is represented by berton & Frey, 1984; Vossler & Pemberton, 1988;
low-diversity, low-density assemblages comprising Beynon & Pemberton, 1992; Pemberton et al., 2001).
Palaeophycus. Other associated ichnofossils include The overall increase in ichnotaxonomic diversity
Diplocraterion, Planolites and ?Conichnus (Fig. 7). In of this trace fossil association relative to that of the
addition, numerous beds are characterised by a bi- Planolites ichnoassociation reflects a fundamental
oturbated texture, in which very few discrete trace change in a number of environmental parameters
fossils can be identified; this could have been above such as nutrient supply, sediment consistency, de-
120 Sunny C. Ezeh, Wilfred A. Mode, Berti M. Ozumba

Fig. 7. Palaeophycus ichnoassociation (well 2; depth 2,496 m). A, B – ?Conichnus (Co) and Palaeophycus (Pa) within sandy
heterolithic facies. Note low diversity and low density of the association; C – Palaeophycus (Pa) and Diplocraterion
(Dp) on heterolithic facies (well 2; depth 2,497 m).

Fig. 8. Asterosoma ichnoassociation (well 1; depth 2,954 m). A, B – Thalassinoides (Th) and Bergaueria (Be) within sandy
heterolithic facies which consists of Asterosoma (As); C (well 1; depth 2,965 m) and D (well 1; depth 2,955 m) illustrate
Asterosoma (As) and escape structure (Es) on sandy heterolithic facies. Scale bar in centimetres.
 Characteristic trace fossils from Miocene brackish-water deposits in the Niger Delta, Nigeria 121

gree of oxygenation and salinity. The end effect was aeoenvironment of the Sorthat Formation on Born-
the establishment of physical and biological condi- holm (Denmark) by Bromley & Uchman (2003). In
tions that were more conducive to benthic coloni- addition, ichnofaunal assemblages of this kind have
sation. been interpreted as characterised by a mixed Cruz-
iana and Skolithos ichnofacies (Howard & Frey, 1973,
5.1.5. Asterosoma ichnoassociation 1975, 1985; Dorjes, 1977; Ekdale et al., 1984, Frey &
Interlaminated mudstones and heteroliths are Pemberton, 1985; Wightman et al., 1987; Beynon &
typified by low-diversity trace fossil assemblages Pemberton, 1992; Tognoli & Netto, 2003).
dominated by reduced forms of Asterosoma (Fig.
8). Associated trace fossil genera include Planolites, 5.1.6. Thalassinoides ichnoassociation
Thalassinoides and Bergaueria. This ichnoassociation This ichnoassociation is monospecific in nature,
is characterised by a predominance of horizontal de- consisting of Thalassinoides (Fig. 9). The illustrated
posit-feeding structures. Although diversity is low, examples show thin laminae or lenses of sands in
individuals, particularly Asterosoma, attain very a matrix of brown mudstones. The mudstones are
high densities, being commonly represented by a preserved in between coarse-grained estuarine
wispy, bioturbated texture in which distinct, other channel sand bodies. These mudstones may repre-
burrow elements are not always readily apparent. sent stable cohesive sediments that were laid down
The ichnogenus Asterosoma is regarded as typical in a protected, low-energy setting that came into ex-
of the transition between the lower shoreface and istence when the estuarine channel was abandoned.
offshore (Ezeh et al., 2016a), but the low degree of The trace fossil characteristic of this ichnoassocia-
bioturbation, low diversity and sediment character- tion has considerable implications; despite the low
istics suggest an environment influenced by salini- diversity and monotypical nature, which is indica-
ty fluctuation, probably a tidally influenced delta. tive of a brackish-water environment, the palaeoen-
This is consistent with the interpretation of the pal- vironment had normal salinity.

Fig. 9. Thalassinoides ichnoassociation (well 3; depth 4,097 m). A – Thalassinoides (Th); B, C – Thalassinoides (Th) on a
brownish mudstone deposit; infill comprises pebbly grained sandstone matrix probably formed during surface
exposure or originating from the overlying sandstone deposit.
122 Sunny C. Ezeh, Wilfred A. Mode, Berti M. Ozumba

5.2. Palaeoenvironmental implications may potentially be utilized as a palaeo-indicator of

brackish-water conditions (compare Frey & How-
Even though the recurring ichnofaunal assemblages ard, 1975, 1980, 1985; Ekdale et al., 1984; Wightman
have significant palaeoenvironmental implications, et al., 1987; Tognoli & Netto, 2003).
the overall trace fossil suite is characterised by: (1) In addition to a decrease in species diversity,
a predominance of morphologically simple, verti- benthic organisms which inhabit brackish-water
cal and horizontal structures; (2) a generally low to environments typically display a reduction in size
very low diversity; (3) reduced size, compared to relative to their normal marine counterparts. For in-
marine counterparts; (4) an association of siderite stance, this is seen in the Asterosoma ichnoassociation
concretions and (5) an admixture of elements which (Fig. 8) where the nominate ichnogenus is smaller
are common to both the Cruziana and Skolithos ich- in form in comparison to the fully marine counter-
nofacies. The overall low ichnotaxonomic diversi- parts recognised by Ezeh et al. (2016a) in the same
ty which typifies the majority of the recurring ich- basin. Similarly, Thalassinoides (Fig. 8) associated in
noassociations differentiated in the Greater Ughelli, the Asterosoma ichnoassociation is typically reduced
Central Swamp and Coastal Swamp depobelts par- when compared to larger forms of Thalassinoides
allels diversity trends documented from modern within the Thalassinoides ichnoassociation which is
brackish-water environments. Ichnoassemblages indicative of fuller marine conditions (Fig. 9). This
in such settings typically are reduced with respect trend is not apparent in non-marine species (Barnes,
to species diversity in comparison to freshwater 1984) which have the ability to adapt to low-salinity
and fully marine counterparts (Beynon & Pember- conditions (Remane & Schlieper, 1971). The relative
ton, 1992; Tognoli & Netto, 2003). This is evident in reduction in size is an adaptive, morphological re-
trace fossil assemblages here assigned to the Gyro- sponse evolved by predominantly marine organ-
lithes-Chondrites, Skolithos and Asterosoma ichnoasso- isms in order to tolerate the high, salinity-induced
ciations. This low diversity is a reflection of the physical and chemical stresses which the inhabited
limited number of benthic species which evolved brackish-water settings impose upon such organ-
physiological specialisations needed to inhabit isms. And, to buttress this point, Remane & Schliep-
brackish-water environments (Barnes, 1984). Very er (1971) argued that reduced salinity affected the
few freshwater species are capable of withstanding size of benthic organisms in a number of ways, in-
salinities in excess of 35 ppm; similarly, very few cluding decreased metabolism, retarded growth
marine species can tolerate salinities lower than 18 and development, promotion of an early onset of
ppm. These species are known as euryhaline organ- sexual maturity, among others, and that the rigours
isms; good examples are the green crab (Carcinus of such waters impose an increased demand for oxy-
maenas) and the green sea urchin (Strongylocentrotus gen on benthic organisms. By decreasing their effec-
droebachiensis). tive surface area these organisms can decrease their
The non-marine and normal marine faunas de- total oxygen consumption and therefore function
note stable end-members at opposite ends of a salin- more efficiently. This reduction in size also serves
ity gradient. With even a slight increase in salinity, as an adaptation to facilitate the osmo-regulation of
the diversity of the non-marine faunal component internal body chemistry due to salinity fluctuations.
declines rapidly. On the other hand, with decreas- The relative morphological simplicity of the
ing salinity the diversity of the normal marine com- trace fossils such as the ones illustrated in Figures
ponent declines at a more gradual rate. Thus, the 4 to 9 reflects the non-specialised feeding strategies
brackish-water faunal assemblage more appropri- employed by the trace-making community. Such
ately represents an impoverished marine assem- organisms are opportunistic in nature and display
blage rather than a true mixture of non-marine and r-selected strategies in population dynamics. Op-
normal marine components (Ekdale et al., 1984; portunistic organisms flourish in areas of high phys-
Wightman et al., 1987). This is illustrated very well iological stress, such as brackish-water settings,
in the Palaeophycus ichnoassociation. Such diversity where animal communities are not resource limited
trends in the distribution of modern benthic organ- (Levington, 1970; Grassle & Grassle, 1974). Oppor-
isms are reflected to some extent in the ancient sed- tunistic or r-selected organisms are characterised by
imentary record by the occurrence of low-diversity rapid reproduction and growth rates, small body
to monospecific ichnofossil assemblages, depicted size, short life cycles, broad environmental toleranc-
by the Planolites and Thalassinoides ichnoassocia- es and non-specialised feeding strategies (Leving-
tions. Various authors have pointed out that such ton, 1970; Jones, 1981; Ekdale, 1985). Such organisms
characteristically low levels of ichnotaxonomic are particularly well adapted to high stress and/or
diversity reflect harsh ecological parameters and low-resource environments (Ekdale, 1985).
 Characteristic trace fossils from Miocene brackish-water deposits in the Niger Delta, Nigeria 123

6. Conclusions References

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