Depth estimation using diameter-thickness ratios in larger

benthic foraminifera
WILLEM RENEMA

Renema, W. 2005 06 15: Depth estimation using diameter-thickness ratios in larger

benthic foraminifera. Lethaia, Vol. 38, pp. 137–141. Oslo. ISSN 0024-1164.

The diameter-thickness ratio (D/T) of larger benthic foraminifera varies indirectly with
depth. Increased turbulence thickens the test, whilst decreased light intensity causes a
flatter test. It is shown that in a nearshore–offshore transect the D/T gradient is similar
between reefs, but stretched over a larger depth gradient. In facies analysis an increase in
D/T can either be caused by decreased hydrodynamic energy, increased transparency or
increased depth (or a combination of these). & Benthic foraminifera, depth, morphometry.

Willem Renema [Renema@naturalis.nl], Nationaal Natuurhistorisch Museum, PO Box

9517, 2300 RA Leiden, The Netherlands; 7th September 2004, revised 7th March 2005.

The morphology of larger benthic foraminifera has
been used for depth-estimation in geological facies
interpretations, usually based on analogues with
recent occurrences. The diameter/thickness ratio
within a species tends to be smaller in shallow
conditions than in deep conditions (e.g. Larsen &
Drooger 1977; Hallock 1979; Hallock et al. 1991). This
tendency was found in symbiont-bearing foraminifera
as a group, among members of the genus Amphistegina
and among individual species in the genus Amphiste-
gina (Larsen & Drooger 1977; Hallock 1979). The
same tendency was found in the Red Sea, the Carib-
bean and Indo-Pacific bioprovinces. Similar results
were found for Palaeonummulites (East Kalimantan;
Renema, unpublished data). In Operculina ammo-
noides a slightly different pattern was found in the Gulf
of Aqaba: specimens at 30 m (the shallowest
measured) were thickest, but the thinnest specimens
occurred at 70–80 m depth, after which thickness
slightly increased (Pecheux 1995). In this study
measurements were taken on dead specimens, and the
decrease in D/T might have been related to tapho-
nomic processes. Pecheux (1995) also found a larger
difference between populations on firm and soft
substrate than with depth.
Experimental work has shown light levels and water
motion influence test thickness (Hallock 1981;
Hallock et al. 1986). In low light levels the test wall
tends to become thinner, whilst in high energetic,
turbulent water the test wall becomes thicker. Gener-
ally, deep settings have lower hydrodynamic energy
levels because they are below storm- and fair-weather
wavebase and experience less tidal-currents. The
penetration of light decreases exponentially with
increased water depth. Thus, shape-change is only
indirectly correlated with depth.
In Amphistegina lessonii the apertural area is orna-
mented with pustules that make a firmer atachment to
the substrate by the pseudopodia possible. In thick-
shelled specimens the apertural area is much larger
than in thin-shelled specimens.
The aim of the present study is to show that shape
trends are influenced by other parameters than water
depth alone. This will limit the use of shape trends for
depth estimation in the fossil record.
Methods
In the Spermonde Archipelago, offshore southwest
Sulawesi (Indonesia) (Figs 1, 2) living larger benthic
foraminifera were collected at reefs at varying differ-
ence from the coast (Renema et al. 2001; Renema &
Troelstra 2001). The D/T ratio of Amphistegina
radiata, A. lessonii and Operculina ammonoides (Figs 3,
4) at four reefs (Lae Lae, Samalona, Kudingareng Keke
and Pulau Lumulumu) at increasing distance from the
shore were analysed. Amphistegina radiata had a
preference for rubble substrate and has been found at
1–6 m water depth nearshore, and 3–33 m offshore.
Amphistegina lessonii had no clear substrate prefer-
ence, and was found on both rubble and sand at 3–9 m
water depth near shore, and 2–36 m water depth
offshore. Operculina ammonoides occurred at the reef
base on both the east and west side of the reefs on sand
substrate. At Kudingareng Keke the sample for
O. ammonoides set was augmented by two grab
samples at 36 and 40 m depth. Only samples from
the west side of the islands were used. During the
southwest monsoon this is the side most exposed to
high, oceanic waves, and the side at which coral
cover is highest. All specimens measured were alive

DOI 10.1080/00241160510013259 # 2005 Taylor & Francis Group Ltd

138 W. Renema
Fig. 1. Map of the Spermonde Archipelago with the sampling sites shown.
at the time of sampling. From each sample 20
specimens were randomly selected using a random
number chart, and diameter and thickness were
measured.
Results
The D/T values found in this study compare very
well with those in previous studies (Tables 1, 2). For
all species there is a good correlation between D/T
and depth at each individual reef (Fig. 5A, B). At
a closer look, however, the D/T ratio is similar
between, respectively, shallow samples in the near
shore reef and mid-slope samples at offshore reefs.
In the shallow samples the thickest tests in both
Fig. 2. Schematic section through a typical reef in the Spermonde Archipelago. The deepest coral cover at the western (seaward) side is 6 m at
Laelae, 18 m at Samalona, 24 m at Kudingareng Keke, and 27 m at Pulau Lumulumu.
LETHAIA 38 (2005)
A. radiata and A. lessonii have been found in the
offshore reefs, and the average D/T decreases towards
the near shore.
The deepest samples of A. lessonii were collected on
soft substrate, compared to firm substrate in the
shallower samples. The D/T ratios in these samples are
slightly higher than those on solid substrate, but the
number of samples is too low to be statistically
significant.
In Operculina ammonoides this pattern is even more
apparent (Fig. 5C). In the shallowest sample D/T is
around 4.0, while in the deepest samples D/T = 6–6.5.
An exception forms the deepest sample at Kudingar-
eng Keke, in which D/T = 5.3. In this sample many
deformed specimens with irregular coiling occur
(Fig. 3).
LETHAIA 38 (2005)
Fig. 3. Habitus of &A. Amphistegina radiata, &B. A. lessonii, &C. Regularly coiled Operculina ammonoides, and &D. Irregularly coiled
O. ammonoides. Scale bars represent 0.5 mm.
Discussion & conclusion
A similar trend with water depth in D/T ratio was
found in three species of larger benthic foraminifera.
Going offshore, water transparency and wave energy
increased (Renema & Troelstra 2001). At each depth,
the near shore samples have a higher D/T than the
more offshore reefs. Thus, generally speaking, the
Fig. 4. Variation in the shape in lateral view of the three species in
this study.
Depth estimation using D/T in foraminifera 139
order (from high to low) of D/T at a given depth is
Laelae–Samalona–Kudingareng Keke–Pulau Lumu-
lumu in the Spermonde Archipelago in the shallow
samples. Similarly, if D/T = 2.8 in Amphistegina
radiata, this would imply a water depth of around
25 m in Palau Lumulumu, but only 10 m in Laelae.
The irregularly coiled specimens of Operculina
occur towards the lower limit of its depth distribution.
In grab-samples taken at 42 and 44 m no living
Operculina were found. In other areas such as the
Berau carbonate shelf just north of the Mankalihat
Peninsula (East-Kalimantan) and South Japan the
maximum depth at which O. ammonoides was found
was 60–70 m water depth (Hohenegger et al. 2000;
Renema, unpublished data). The relatively shallow
lower depth limit of larger benthic foraminifera in the
Spermonde Archipelago is caused by seasonal varia-
tion in transparency (Renema & Troelstra 2001). The
irregular growth of Operculina ammonoides could be
caused by sub-optimal conditions during the wet
monsoon.
These results show that D/T ratios cannot directly
be interpreted towards depth and differ between
species. This restricts the use of the D/T ratio for depth
estimation in facies interpretation. An increase in D/T
can either mean a decrease in hydrodynamic energy,
140 W. Renema LETHAIA 38 (2005)

Table 1. D/T measurements previously reported.

Species D/T Depth Exposure Region Source

Amphistegina spp 1.99 10 Seaward Palau Hallock 1979

Amphistegina spp 2.03 20 Seaward Palau Hallock 1979
Amphistegina spp 2.08 2 Lagoonal Palau Hallock 1979
Amphistegina spp 2.14 20 Lagoonal Palau Hallock 1979
Amphistegina spp 1.78 1 Seaward Hawaii Hallock 1979
Amphistegina spp 2.04 30 Seaward Hawaii Hallock 1979
A. lobifera 1.72 0–10 Hallock 1979
A. radiata 2.5 15–30 Hallock 1979
A. bicirculata 2.4 30–100 Hallock 1979
Heterostegina depressa 4 5–50 Hallock 1979
Operculina ammonoides 5.5 20–100 Hallock 1979
A. lobifera 1.57–1.79 5 Gulf of Elat Hallock & Hansen 1979
A. lobifera 1.88–2.06 45 Gulf of Elat Hallock & Hansen 1979
A. lessonii 1.98–2.07 5 Gulf of Elat Hallock & Hansen 1979
A. lessonii 2.17–2.29 38 Gulf of Elat Hallock & Hansen 1979
A. lessonii 2.24–2.42 64 Gulf of Elat Hallock & Hansen 1979
A. papillosa 2.27–2.60 45 Gulf of Elat Hallock & Hansen 1979
A. papillosa 2.87–3.13 70 Gulf of Elat Hallock & Hansen 1979
A. papillosa 3.65–4.37 90 Gulf of Elat Hallock & Hansen 1979
A. bicirculata 2.84–3.41 45 Gulf of Elat Hallock & Hansen 1979
Operculina ammonoides 3.7–5.3 40 Soft substrate Pecheux 1995
O. ammonoides 1.9–4 40 Hard substrate Pecheux 1995
O. ammonoides 5.5–11 85 Soft substrate Pecheux 1995
O. ammonoides 3–9 130 Soft substrate Pecheux 1995

Table 2. D/T measurements (average and standard deviation) for each sample of 20 specimens.

Amphistegina lessonii A. radiata Operculina ammonoides

Island Substrate Depth D/T SD D/T SD D/T SD

Laelae 3 2.38 0.06 1.97 0.25

6 2.51 0.08 2.48 0.23
9 3.01 0.07
Samalona 3 2.27 0.10 1.60 0.21
6 2.43 0.09 2.05 0.29
9 2.52 0.08 2.55 0.23
12 2.62 0.07 2.64 0.18
15 2.59 0.12 2.88 0.31
18 2.76 0.10 3.9 0.30
21 2.87 0.12 4.87 0.34
24 6.16 0.60
Kudingkarengkeke 3 2.24 0.08 1.74 0.15
6 2.37 0.10 1.83 0.21
9 2.41 0.09 1.95 0.22
12 2.57 0.12 2.35 0.18
15 2.64 0.05 2.28 0.15
18 2.70 0.08 2.5 0.17
21 2.74 0.07 2.72 0.25
24 2.75 0.12 2.99 0.27 4.12 0.25
sand 27 2.85 0.09 4.53 0.28
sand 30 2.92 0.13 5.35 0.25
sand 33 5.80 0.32
sand 36 6.0 0.29
sand 40 5.34 0.70
Pualu Lumulumu 3 2.18 0.08 1.33 0.23
6 2.23 0.10 1.60 0.34
9 2.44 0.09 2.17 0.26
12 2.5 0.11 2.45 0.24
15 2.55 0.12 2.56 0.23
LETHAIA 38 (2005) Depth estimation using D/T in foraminifera 141

Table 2. Continued.
Amphistegina lessonii A. radiata Operculina ammonoides

Island Substrate Depth D/T SD D/T SD D/T SD

18 2.59 0.10 2.35 0.25

21 2.66 0.09 2.56 0.23
24 2.64 0.06 2.73 0.18
27 2.70 0.12 2.95 0.26
sand 30 2.79 0.13 4.18 0.23
sand 33 2.82 0.10 4.93 0.18
sand 36 5.66 0.26
sand 39 6.25 0.32

an increase in transparency or deepening. By using

only D/T data it is not possible to discriminate
between these causes.
Assemblage composition can help in determining
the location in the onshore–offshore gradient in
modern settings. In near shore soft bottom assemblage
the only nummulitid species is O. ammonoides, while
going offshore sequentially Palaeonummulites venosus
and Operculina complanata occur in this assemblage as
well (Renema 2002). However, these gradients are not
well documented in fossil carbonate shelves yet, and it
is difficult to infer the position in the near shore–open
sea gradient in fossil settings.

Fig. 5. Depth vs diameter/thickness (D/T) in &A. Amphistegina
radiata, &B. A. lessonii and &C. Operculina ammonoides.
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