Benthic Foraminifera as Environmental Indicators

in Torres Strait–Gulf of Papua

*Alexandra L. Post, Laura Sbaffi, Vicki Passlow and Dylan C. Collins
Geoscience Australia, GPO Box 378, Canberra, ACT 2601, Australia
*Corresponding author, E-mail: Alix.Post@ga.gov.au

Post, A.L., Sbaffi, L., Passlow, V., and Collins, D.C., 2007, Benthic foraminifera as environmental indicators in Torres
Strait–Gulf of Papua, in Todd, B.J., and Greene, H.G., eds., Mapping the Seafloor for Habitat Characterization:
Geological Association of Canada, Special Paper 47, p. 329-347.

Abstract

Three areas in the Torres Strait–Gulf of Papua region were selected for detailed study of sediments and benthic fossil
biota. These areas form a transect across the shelf from the Fly River Delta to the shelf edge, near the northern extrem-
ity of the Great Barrier Reef. The Torres Strait–Gulf of Papua shelf is a shallow, low-gradient platform, where the shelf
edge occurs between 120 and 140 m depth. In the study area, where the sediments range from muddy to gravelly car-
bonate sands, the sediment deposition rates are low and the relict content of sediment is often high.

The three areas show distinct differences in benthic foraminiferal assemblages as indicated by relative abundances at
the order level, as well as distribution patterns of individual species; these differences are also reflected in the total
microbiotic communities. Given the high relict content in the surface material across these sites, a foraminiferal preser-
vation scale was developed to assess the extent of reworking. Taphonomic features indicate that abrasion is the main
factor affecting preservation.

Despite poor preservation of the foraminiferal tests, the benthic foraminiferal species have a strong correlation to water
depth, indicating that transportation pathways are short. Application of multivariate statistics to analyze the relation-
ship between environmental attributes and the distributions of the microbiota and foraminiferal species indicates the
additional importance of factors including percent carbonate mud, percent gravel, organic carbon flux, temperature,
salinity and mean grain size. The benthic foraminifera produce a much stronger correlation to the environmental vari-
ables than the microbiota, indicating that these organisms can provide a detailed assessment of habitat types.

Résumé

Trois zones dans le détroit de Torres, golfe de la région de la Papouasie, ont été sélectionnées pour l'étude détaillée des
sédiments et du biote fossile benthique. Ces zones forment un transect dans la plate-forme du delta du fleuve Fly jusqu'à
l'accore du plateau, près de l'extrémité nord de la Grande Barrière de corail. Le détroit de Torres dans le plateau du
golfe de la Papouasie est une plate-forme de faible gradient, où l'accore du plateau se trouve entre 120 et 140 m de pro-

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POST ET AL.

fondeur. Dans la région à l'étude, où les sédiments varient de sables carbonatés boueux à graveleux, les taux de sédi-
mentation sont faibles et le contenu en reliques des sédiments est souvent élevé.

Les trois zones montrent des différences dans les assemblages de foraminifères benthiques comme l'indiquent les abon-
dances relatives au niveau de l'ordre, ainsi que les patrons de distribution des espèces individuelles; ces différences se
reflètent aussi dans l'ensemble des communautés microbiotiques. Étant donné le fort contenu en reliques du matériel de
surface dans ces sites, une échelle de préservation des foraminifères a été définie pour évaluer l'étendue du
remaniement. Les caractéristiques taphonomiques indiquent que l'abrasion est le principal facteur qui affecte la préser-
vation.

Malgré une faible préservation des tests de foraminifères, les espèces de foraminifères benthiques montrent une forte
corrélation avec la profondeur de l'eau, ce qui indique que les voies de transport sont courtes. L'application de statis-
tiques à variables multiples dans l'analyse des relations entre les caractéristiques environnementales et les distributions
des espèces du microbiote et de foraminifères montre l'importance additionnelle de facteurs comme le pourcentage de
boue carbonatée, le pourcentage de gravier, le flux de carbone organique, la température, la salinité et la taille moyenne
des grains. Les foraminifères benthiques montrent une corrélation beaucoup plus grande avec les variables environ-
nementales que le microbiote, ce qui indique que ces organismes peuvent mener à une évaluation détaillée des types
d'habitats.

INTRODUCTION
Analysis of the relationships between marine organisms and their
environment is becoming an important tool for understanding and
managing marine communities (Hockey and Branch, 1997;
Williams and Bax, 2001; Roff et al., 2003). Benthic foraminifera
are a group of organisms that are ubiquitous throughout the world’s
oceans, but the distribution of individual species is constrained by
their environmental preferences. Therefore, these organisms have
the potential to provide important environmental information.
Benthic foraminiferal species distributions may vary according to
any combination of factors such as substrate type, light intensity,
water temperature, food availability, oxygen, salinity and current
energy (Murray, 1991). Many of these attributes vary with water
depth, so depth may capture variations in these parameters and in
species distributions. Understanding the response of the whole
ecosystem to environmental change is important for marine man-
agement. This study tests the environmental controls on the distri-
bution and abundance of benthic foraminiferal species and other
microbiota preserved in surface sediments as a means of character-
izing habitats. The environmental-biota relationships determined in
this study will also allow interpretation of environmental and
ecosystem change through time, when applied to down-core records
of cores collected from this region.
Benthic habitat mapping from offshore Canada and New
Zealand illustrates the success of defining biological communities
based on the nature of the substrate as well as water depth (e.g.,
Thouzeau et al., 1991; Kostylev et al., 2001; Hayward et al., 2002).
However, studies from other regions, including northern Australia,
reveal the difficulties of relating biological and habitat data in
regions of slow sedimentation with strong reworking (e.g., Orpin et
al., 1999). In these areas of high reworking, the relationship
between sedimentary biotic material and physical parameters, such
as the substrate type and water depth, is degraded, making it diffi-
cult to define key habitat attributes. The use of sedimentary materi-
al to delineate habitats and benthic communities may therefore fail
in high-energy regions.
In this study, surface sediments from across the Torres Strait–
Gulf of Papua region are analyzed to assess the relationship
between environmental attributes and the distribution of benthic
foraminiferal species and other microbiota. The Torres Strait area is
a very dynamic region, encompassing sub-delta to open marine
environments. The dynamic nature of this region provides a good
basis for testing the relationship between the sedimentary biotic
record and physical features of the environment. Further, this study
incorporates taphonomic data to determine the extent to which
reworking can alter the relationship between biota and environmen-
tal attributes. Sediment transport, sediment mixing and test dissolu-
tion have been found to alter the composition of foraminiferal
assemblages from the Fly River Delta (Torres Strait; Cole, 1995).
SETTING
Sampling was conducted across three main areas: Areas A, B and C
(Figure 1). Area A is located at the distal edge of the Fly River Delta
within 30-40 km of the shoreline and at water depths ranging
between 25 and 35 m. The area is mainly flat to undulating, has low
relief ridges (up to 2 m), and swales also occur parallel to the bathy-
metric contours. A shallow gully (~3 m) occurs in the southwest and
trends northwest–southeast. Terrigenous sediment transport and
accumulation rates are low across this area as indicated by the thin
Holocene section present in cores (typically <1 m thick) and the
abundance of semi-lithified mud clasts (Harris et al., 2004). Many
sites exhibit a bimodal grain size distribution reflecting the accumu-
lation of mud from the tidal flats and from the reworking of fine
sediments.
Area B is located 75 to 85 km offshore and contains a subma-
rine valley system that trends east–west across the shelf in two sep-
arate limbs (Figure 1). The northern valley is interpreted as a river/
estuarine channel, while the southern system is characterized as a
tidal channel (Heap et al., 2003). The valley reaches depths of up to
130 m, with an overall depth across the area of between 40 and 130
m. Shallower areas include shoal reefs that occur in the southern-
most section of Area B. Shallow gullies (~10 m) are also found, and

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Figure 1. Grab samples were collected at 64 sites within areas A, B
and C in the Torres Strait–Gulf of Papua region, southeast of the
Fly River Delta.
these drain into the valley system. The areas that were sampled
were generally flat and included some rocky and reef environments.
Area C is located 105 to 120 km offshore and comprises a sub-
marine valley system aligned west-northwest–east-northeast
(Figure 1). The valley occurs as two separate limbs with the deeper
southern channels representing relict features formed by tidal cur-
rent scour during Pleistocene sea level lowstands (Heap et al.,
2003). The water depth at sampling sites across this area ranges
from 50 to 220 m, and the shallow areas are mapped as platform
reefs. The channels across both areas B and C act as conduits for
upwelled Coral Sea water, which is relatively cool, saline and nutri-
ent-rich (Harris et al., 2004).
Bottom measurements of temperature and salinity at each site
indicate that each of these parameters is tightly correlated with
changes in water depth. As water depth increases, the temperature of
the bottom water decreases logarithmically (R2 = 0.75) while salini-
ty increases (R2 = 0.80). The bottom water temperature ranges
between 25 and 29°C across all sites, and salinity between 34.33 and
35.18‰. The increase in salinity with depth is associated with
upwelling of saline, cool water from the Coral Sea into deeper chan-
nels of areas B and C (Harris et al., 2004), while shallower nearshore
areas are influenced by fresh water input from the Fly River.
METHODS
Analysis of Environmental Parameters
Surface sediment samples were collected from 64 sites within the
Torres Strait–Gulf of Papua region during Geoscience Australia
BENTHIC FORAMS AS ENVIRONMENT INDICATORS
Survey 234 aboard RV Franklin in January-February 2002 (Figure
1). The physical characteristics of each site were determined
through several analyses. Surface grab samples were analyzed for
gravel, sand and silt content by washing through 2 mm and 63 μm
sieves, and the <2 mm fraction was analyzed by laser particle size.
Carbonate content was determined on the mud and sand fractions
using the carbonate bomb method. In addition, the seabed morphol-
ogy was characterized based on swath bathymetry data, together
with seabed photography. Temperature and salinity measurements
were obtained at each site via conductivity/temperature/depth
(CTD) casts. The measurements from nearest the seabed were used
in comparison with the benthic foraminiferal assemblages. Details
of each of these analyses are available in Harris et al. (2004).
Analysis of Benthic Foraminifera
Surface sediments were collected using a Smith-McIntyre grab. The
grab material was dry-sieved to the 150 :m to 2 mm size fraction
and split into aliquots, each containing at least 300 specimens of
benthic foraminifera. The foraminifera were identified using a
binocular light microscope and referencing the taxonomic studies of
Hottinger et al. (1993), Loeblich and Tappan (1994), Hayward et al.
(1997) and Haig (2002). The most abundant forms (>2% abundance)
were identified to species level (see species list in Appendix I), while
rarer forms were grouped and identified to genera.
Taphonomy
Two foraminiferal genera were selected for assessment of taphono-
my, following an overview of preservation in samples from all three
areas. Both genera selected: Amphistegina sp. and Quinqueloculina
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sp., occur in reasonable abundances across the whole study area and
together encompass robust to fragile forms. The assessment was
restricted to non-agglutinated forms because of the very different
nature of taphonomic breakdown in agglutinated taxa. A preserva-
tion scale was determined for each genera (Table 1), based on fea-
tures documented from field and laboratory studies (Peebles and
Lewis, 1991; Kotler et al., 1992; Shroba, 1993) and the degree to
which they affect the test; typical examples are illustrated in Figure
2. The scale was designed to allow comparisons between genera,
while taking into account the different types of degradation that
occur between species.

Analysis of Microbiota

The absolute abundance of the main groups of organisms identi-

fied in the sedimentological size fraction 150 :m–2 mm was also
calculated for this study. The 64 grab samples were repeatedly
split using a microsplitter and weighed using a Sartorius BP221S
precision scale to obtain about 0.2 gm of sediment. All the shells
and recognizable fragments present in the subsamples were then
counted and expressed as number of specimens/g of dry sediment.
Of the 15 recognized groups, two live in the water column (i.e.,
planktonic foraminifera and pteropods) while the others mainly
dwell on the seafloor and/or a few centimetres within the sedi-
ment. This range of taxa allowed a more complete picture of the
marine ecosystem.
Figure 2. Preservation styles for Quinqueloculina subimpressa (A
DATA ANALYSES and B) and Amphistegina papillosa (C and D). A shows example of
extensive surface pitting, holes and scalloping of test edges, while
Statistical analysis was conducted using PRIMER-E software B shows good surface preservation but a high degree of fragmenta-
(Plymouth Routines In Multivariate Ecological Research; see Clarke tion of the test. C illustrates a specimen with light, mainly surface
and Ainsworth, 1993; Clarke and Warwick, 2001). Similarities alteration, and D shows heavy surface abrasion and fragmentation
between sites were analyzed based on benthic foraminiferal species of the test.
data and environmental attributes to assess inter-relationships.

Table 1. Preservation scales for Amphistegina sp. and Quiqueloculina sp.

Taphonomic Features
Scale Amphistegina sp. Quinqueloculina sp.

Fresh Surface transparent, details of pores clearly visible; Surface transparent to translucent, pitting and breakage
appearance breakage minor <10% (usually scalloping) minor, <10%

Lightly Surface transparent to translucent; minor breakage or Surface translucent to opaque; minor damage <25% of test
modified scalloping <25% of test; surface ornament minor change edge scalloped or broken or <25% breakage; surface shiny
(e.g., ornament smoothing or removal, pitting) or ornament shows minor change

Moderately Surface transparent to opaque; <50% of edge scalloped Surface opaque ± minor staining; up to 50% of edge
modified or <a of test broken; <50% of surface affected by scalloped and/or up to a of test broken; surface alter-
holes, pitting, ornament smoothing or surface removal ation on <50% of test surface (e.g., ornament smoothing,
pitting, surface removal)

Heavily Surface opaque, surface ornament smoothed or surface Surface opaque, ornament smoothed or surface layers
modified layers partially removed, >50%; breakage if present, partially removed, >50%; test may be heavily stained;
>30%; infill if present not lithified breakage and/or abrasion and pitting >30%

Relict Surface ornament smoothed or layers removed or test Surface ornament smoothed or polished, or surface sugary,
polished or heavily etched; staining, if present, heavy >75% affected and/or test heavily stained, >75% of area;
>75%; breakage usually >50% of edge or whole test; test breakage >50% of test or edge; test infilled and/or lithified
infilled and lithified

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 BENTHIC FORAMS AS ENVIRONMENT INDICATORS

Analysis of Community Structure
Similarity between sites was determined based on counts of the
microbiota (measured as the number of individuals/gram of sedi-
ment) using the Bray–Curtis similarity coefficient. The Bray–Curtis
similarity coefficient provides good results for ecological data
because it does not measure the absence of species (Clarke and
Ainsworth, 1993). This is important for biological studies because
a species may be absent at either end of its ecological range. For the
benthic foraminifera, species counts were square-root transformed
to allow stronger weighting of rarer forms, and only species that
have abundance >2% were used. Microbiota were identified to phy-
lum, and these counts were also square-root–transformed. Principal
Components Analysis (PCA) was used to obtain a broad indication
of the species which account for the greatest variance in the dataset.
Analysis of Environmental Parameters
The similarity between sites based on the microbiota abun-
dances is shown as a Multi-Dimensional Scale (MDS) plot in
Figure 4. MDS plots depict the relative differences between sites by
the distance between points on the plot (Kruskal and Wish, 1978).
The site similarity shown in Figure 4 indicates that samples from
Area A form the most distinct grouping. Considerable overlap in
microbiota abundances is observed within areas B and C. The over-
lap between areas B and C reflects overall similarities in the abun-
dances of the benthic foraminifera (45 ± 10% in both areas) and of
several of the other dominant phylum (e.g., pteropods 14% and bry-
ozoan 7-8%).
The dominance by the benthic foraminifera across each of the
sites highlights the importance of this group in the region and the
Table 2. Environmental attributes and their transformation for
Torres Strait surface sediments

Similarity between sites was determined for the environmental Environmental attribute Transformation
parameters based on normalized Euclidean distance. The key envi-
ronmental attributes and their transformations are listed in Table 2. %Mud Log
Several attributes were log-transformed to reduce the skewness in %Sand None
their distribution across sites, allowing a more meaningful normal- %Gravel Log
ization of the data. Another parameter which was included in the Mean grain size None
analysis was the number of benthic foraminifera/g of sand (BF/g). %CaCO3 (total) Log
The concentration of benthic foraminifera in the sediment is con- %CaCO3 mud Log
trolled by the flux of organic carbon to the seafloor at depths above %Terrigenous mud Log
the lysocline (see Herguera, 1992). For this study the number of Water depth (m) Log
benthic foraminifera was calculated in a weighed aliquot of sedi- BF/g None
ment in the 150 :m–2 mm size fraction, and this was applied as a Temperature (°C) None
proxy for organic carbon flux. Salinity None

Comparison between Distribution of Biota and

Table 3. Average abundances across each area for the 15 groups of
Environmental Attributes marine organisms recognized and counted in the Torres Strait grab
samples
The BIO-ENV function in PRIMER was used to determine the
strength of the relationship between the biota and measured envi- Percent relative abundance
ronmental variables (Clarke and Ainsworth, 1993; Clarke and ± standard deviation
Warwick, 2001). The environmental and biotic matrices were com- Faunal Group Area A Area B Area C
pared using Spearman’s rank correlation, which is appropriate for
comparing similarity matrices derived from different types of coef- Serpulids 0.05 ± 0.05 0.1 ± 0.2 1.0 ± 0.9
ficients (i.e., the Bray–Curtis coefficient for the species matrix and Coralline algae 0.1 ± 0.2 0.4 ± 0.3 0.6 ± 0.7
Euclidean distance for the environmental matrix). The BIO-ENV (Halimeda) fr
function proceeds by taking each environmental matrix in turn to Corals fr 0.3 ± 0.4 0.4 ± 0.4 0.6 ± 0.5
determine which combination of factors best represents the species Brachiopods 0.3 ± 0.2 0.6 ± 0.5 0.7 ± 0.3
distribution. Porifera fr 0.5 ± 0.3 0.5 ± 0.5 0.6 ± 0.4
Planktonic 0.8 ± 0.2 5.8 ± 3.1 10.6 ± 5.2
RESULTS foraminifera
Echinoderms fr 1.1 ± 1.3 2.2 ± 0.9 2.0 ± 0.9
Abundance of Total Microbiota Soft corals fr 1.3 ± 3.3 4.0 ± 3.0 3.6 ± 1.4
Gastropods 2.3 ± 1.0 3.1 ± 1.1 3.3 ± 1.5
Benthic foraminifera are the dominant phylum in terms of their rel- Ostracods 2.5 ± 0.6 2.0 ± 0.9 2.5 ± 1.3
ative abundance in sediments across all areas, with abundances of Other arthropods fr 3.0 ± 3.0 3.0 ± 1.8 3.0 ± 1.8
~45% in areas B and C and ~70% in Area A (Table 3, Figure 3). Bryozoa fr 3.2 ± 5.0 7.3 ± 2.2 8.2 ± 4.2
The next most dominant taxa are pteropods (7.5-14.5%), bivalves Pteropods 7.6 ± 4.9 14.4 ± 6.9 13.4 ± 4.8
(5-10%), planktonic foraminifera (1-10.5%) and bryozoa (3-8%). Bivalves 7.7 ± 2.1 9.5 ± 3.8 4.8 ± 2.1
The other phyla have average abundances of less than 5% across Benthic foraminifera 69.3 ± 12.4 46.5 ± 10.8 45.1 ± 9.2
each area.
Note: fr denotes presence of fragments only

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POST ET AL.
Figure 3. Average abundances of microbiota across areas A, B and C. The abundance of benthic foraminifera decreases between areas A
and C, while planktonic foraminifera and bryozoa increase.
Figure 4. Site similarity (MDS) based on the abundance of micro-
biota (measured as number of specimens/gram of sediment).
Similarity is plotted as the relative distance between sites. Sites
within Area A show a distinct difference in microbiota abundances
to areas B and C. There is substantial overlap in site similarities
within areas B and C.
potential for mapping the ecosystem through detailed studies of
species within this phylum.
Foraminiferal Abundances
Benthic foraminifera are abundant in each of the grab samples
examined. Thirty-one species >2% and minor species (numerical)
334
were incorporated within 15 genera. Species abundances were cal-
culated as percentage and the number of foraminifera/g of sediment
in the sand fraction.
Principal components analysis indicates that most of the vari-
ance (68%) within the species data can be explained by the distri-
bution in the abundances of Amphistegina papillosa, Assilina
ammonoides, Cibicides refulgens, Cibicides-type species,
Nummulites venosus, Heterolepa subhaidingeri, Pseudogaudryina
pacifica and Spiroplectinella pseudocarinata. The distribution of
these species exhibits distinct trends with respect to water depth
(Figure 5). Amphistegina papillosa, A. ammonoides and N. veno-
sus are most abundant in waters <60 m deep, and are therefore the
dominant species within Area A. At depths below 60 m, A. papil-
losa reaches average abundances of 30%, while A. ammonoides
and N. venosus have maximum average abundances of 20% and
10%, respectively. Species with maximum abundances at mid-
depths (between 50 and 110 m) include P. pacifica (maximum at
55 m), S. pseudocarinata (maximum at 110 m) and H. subhaidin-
geri (maximum at 85 m). These species tend to increase in abun-
dance across Area B and shallower parts of Area C.
Pseudogaudryina pacifica maintains its presence across the deep-
er water sites and has an abundance of 6%. Cibicides-type species
and C. refulgens are distributed across all water depths and the
highest abundance occurs at depths >120 m for Cibicides-type
species. The maximum abundance of C. refulgens occurs at 90 m.
However, it maintains high abundances across depths >120 m.
These two species are most common within Area C.
The distribution of several of these key species across the sites
is shown in Figures 6A-C, illustrating the relationship between
bathymetry and species abundances on a site-by-site basis. Assilina
 BENTHIC FORAMS AS ENVIRONMENT INDICATORS

Figure 5. Percent abundance of major benthic foraminiferal species according to water depth. Trendlines are shown as moving averages.
Note that the scale changes between plots.

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POST ET AL.
ammonoides (Figure 6A) has consistently high abundances across
Area A, while high abundances of this species within areas B and C
are restricted to the shallower plateaux, and very low abundances
occur at deeper sites. A similar distribution is shown for A. papil-
losa in Figure 6B, with the highest abundance of this species in
areas B and C occurring at the shallowest sites. Spiroplectinella
pseudocarinata (Figure 6C), by contrast, shows highest abundances
at the deepest sites, and therefore has very low abundance at sites in
areas A and B, with slightly increased numbers occurring at the few
relatively deeper sites in Area B.
Cluster analysis of the foraminiferal species data reveals 7
main clusters. Cluster 1 consists of just one sample from Area A,
Site 4, with a similarity to other sites of 50%. Other sites from Area
A form Cluster 2 (Figure 7), with similarities ranging between 70
and 90%. The foraminiferal assemblage at Site 4 contains higher
abundances of Quinqueloculina philippinensis and Cibicides-type
species, and much lower abundances of N. venosus compared to the
assemblages at other sites within Area A. Clusters in areas B and C
show some overlap between the two areas, with Cluster 4 the only
distinct cluster, composed of sites from the deeper parts of Area C
(Figure 7). Similarities between sites within these remaining clus-
ters range between 60-85%.
336
Figure 6. Percentage abundance of A. ammonoides
(A), A. papillosa (B) and S. pseudocarinata (C) at
each site. Species show a clear relationship to depth,
with highest abundances of A. ammoinoides and A.
papillosa in Area A and the shallower plateaux with-
in areas B and C, and highest abundances of S.
pseudocarinata associated with the deeper water
sites in areas B and C.
Taphonomic Features of Benthic Foraminifera
The taphonomic changes in selected genera of benthic foraminifera
have been analyzed across all the sites. All samples show evidence
of some form of mechanical or chemical breakdown, but with vari-
able intensity between sites. Effects observed include pitting, break-
age, scalloping of test edges, flattening or removal of surface orna-
mentation or surface layers, polishing, test infill and staining
(Figure 2). Abrasion features are the most common form of test
alteration, indicating mechanical breakdown due to bombardment
with fine-grained material. Dissolution effects, mainly seen as
opacity of the test surface, are also common. The very heavy abra-
sion of some specimens indicates their long residence time near the
surface. There appears to be no correlation between species abun-
dances and their preservation, implying that preservation is not a
function of species production or transportation to the sites.
The preservation of Amphistegina sp. is poor in Area A, sug-
gesting a high degree of reworking in these sediments. Areas B and
C, however, are dominated by sites with >25% of tests with a fresh
appearance or lightly modified (Figure 8). Across the remaining
sites, most of the tests fall into the heavily modified or relict cate-
gories. Average preservation values of Amphistegina sp. across all

Figure 7. The distribution of species clusters across areas A, B and C. Data is based on the abundance of benthic foraminiferal species
(measured as number of specimens/gram of sediment). Sites within Area A form distinct clusters, with areas B and C showing greater over-
lap in clusters between the two areas.
three areas are shown in Figure 9. The detailed breakdown of
preservation for Amphistegina sp. is provided in Appendix II.
In Area A, samples are generally dominated by tests showing
heavy abrasion, typically >40-70%. The abundance of both relict
and material of fresh appearance is generally low, <10%; 6 samples
have no relict specimens, while 5 of the 21 samples show values
>20%. Similarly, the abundance of material of fresh appearance is
low, 0-10%; 3 samples have no tests of fresh appearance.
Area B shows greater variability. Samples are generally domi-
nated by tests that are heavily degraded, typically in the range >25-
65%. The relict content is variable; 7 samples have no relict speci-
mens, the remainder range from 1-35%. However, the percentage of
material of fresh appearance and light modification is generally
higher than in Area A. Area C also shows a high level of variabili-
ty. About half of the samples are dominated by tests in the heavily
modified category, although the dominance is much less clear cut
than in samples from areas A and B. The remaining samples are
dominated by light or moderate abrasion, or, in one case, by mate-
rial of fresh appearance.
The preservation of Quinqueloculina sp. is poor throughout;
sites with poor preservation outnumber sites where well-preserved
material dominates (Figure 8). Average preservation values across
BENTHIC FORAMS AS ENVIRONMENT INDICATORS
all sites are shown in Figure 10. Details of Quinqueloculina sp.
preservation are provided in Appendix III. In Area A, samples are
dominated by material that has been heavily degraded, >40-78%.
Relict content is low (0-20%) to moderate (>20-45%). The content
of material of fresh appearance is low throughout, <5%, with 12 of
the 21 samples having no specimens. The proportion of lightly
modified material is low, generally in the range 0-20%, with 9 sam-
ples having no lightly modified specimens.
Area B samples are generally dominated by material with
heavy degradation, 30-50%, although some samples show higher
levels of either relict or moderately abraded material. There is
greater variability in the relict content where 14 out of the 21 sam-
ples have low values (0-20%) and the remainder range from 20 to
49%. The proportion of material of fresh appearance is similarly
variable, 0-18%; 14 of 21 samples have no specimens with a fresh
appearance. Where present, the proportion of material of fresh
appearance is generally higher than in Area A samples, as is the pro-
portion of lightly modified material.
In Area C, samples show greater variability; some contain a
higher content (up to 50%) of material that has a fresh appearance
and light modification, with little or no relict material; others are
biased towards being heavily abraded and relict, and contain lit-
tle material with fresh appearance or light modification. Relict
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Figure 8. Preservation of benthic foraminifera for Amphistegina sp. and Quinqueloculina sp. Preservation is shown for sites that have 25-
50% and >50% well-preserved specimens (based on the fresh appearance and lightly degraded categories). There are 12 sites with >50%
well-preserved Amphistegina sp. and 2 with well-preserved Quinqueloculina sp. Best preservation occurs within Area C.

Amphistegina spp.

heavy heavy
heavy

relict
relict relict

fresh
fresh moderate fresh
light
moderate
moderate light light

04-070-9

Area A Area B Area C

Figure 9. Relative preservation, averaged across each area, based on Amphistegina spp. The combined relict and heavily degraded catego-
ry is highest across areas A and B.

338
 BENTHIC FORAMS AS ENVIRONMENT INDICATORS

Quinqueloculina spp.

relict
heavy
relict relict
heavy heavy

fresh
fresh fresh
light

light light
moderate
moderate moderate

04-070-10

Area A Area B Area C

Figure 10. Relative preservation, averaged across each area, based on Quinqueloculina spp. Most sites have a high relict content, with most
material of fresh appearance occurring in Area C.

content varies between 0-60%; 4 samples have no relict material.
Heavily modified content varies from 16-70%. Six samples
have no material of fresh appearance; the remainder varies
between 3-26%. The proportion of lightly modified material has
a similar range with three samples having no lightly modified
material.
Relationship between Biota and
Environmental Variables
remaining clusters in areas B and C, with highest fluxes in Cluster
6. There is no correlation between BF/g and grain size across the
study sites, indicating that the benthic foraminiferal accumulation is
not a reflection of sediment transport. In the following section, the
relationship between the distributions of the environmental vari-
ables shown in Table 4 and the benthic foraminiferal assemblages
and the general biota are examined and tested.
Benthic Foraminifera

Several key environmental attributes vary between the species clus-
ters (Table 4). Mean water depth varies substantially between the 7
clusters, ranging from 24.5 m in Cluster 1 to 119 m in Cluster 4
(Table 4). The grain size and carbonate content also show a large
range in values across the sites. The number of benthic
foraminifera/g (BF/g) indicates a high organic flux in Cluster 2
(Area A), compared to the exceptionally low flux in Cluster 1 (Site
4, also Area A). The organic flux is more moderate across the
Analysis of individual parameters indicates that depth has a strong
correlation to the site similarity expressed by the benthic assem-
blages, with a Spearman’s rank correlation coefficient (Ds) of 0.68
(Figure 11A). Stronger correlations were found when factors were
combined, with the best correlation observed for the combination of
% gravel, % CaCO3 mud, depth, organic carbon flux, temperature
and salinity (Ds = 0.802). However, the combination of just three
factors, depth, organic carbon flux and salinity (DOS), explain near-

Table 4. Average values for environmental variables within each species cluster

Cluster 1 Cluster 2 Cluster 3 Cluster 4 Cluster 5 Cluster 6 Cluster 7

%Mud 3.7 23.4±10.8 15.6±14 8.8±4.8 15.3±5.4 20.5±7.6 2.9±0.4

%Sand 26.7 70.8±10.2 71.3±16 81.6±8 71.9±15.8 68.4±21.1 66.8±1.9
%Gravel 69.6 5.8±4.8 13.1±10.1 9.6±8.1 12.8±11.8 11.1±16.5 30.2±2.3
Mean grain N/A 299±142 277±132 305.9±97 251±47 200±106 623±39
%CaCO3 M 0.4 2.8±0.8 10±10.7 7.3±3.9 10.5±3.1 15.2±6.8 2.2±0.6
%CaCO3 T 92.1 54.4±11.6 62.5±22.2 64.2±19.4 54.4±14.8 72.8±16 93.8±3
Water depth 24.5 31.3±2.0 61.9±14.1 119.2±40.4 76.6±12.9 81.7±28.5 50.0±0
BF/g 395 4574±1403 2680±839 2663±921 1747±602 5176±1448 1727±850
Temperature 28.9 28.7±0.3 27.5±0.2 25.4±0.4 27.2±0.7 27.1±0.9 27.5±0.4
Salinity 34.48 34.50±0.08 34.91±0.06 35.15±0.03 34.97±0.09 34.95±0.13 34.95±0.1

Notes: ‘Mean grain’ refers to the mean grain size <2 mm, '%CaCO3 M' and '%CaCO3 T' refers to percentage carbonate in the mud fraction
and in the total sample respectively, water depth is measured in metres, ‘BF/g’ is the number of benthic foraminifera in each gram of sand.
There was insufficient material in the sand fraction to measure the mean grain size in sample 4 (Cluster 1).

339
POST ET AL.

BioEnv results
A
Mean G rain size
% Sand
%Mud
%Gravel
% CaCO3 Mud
O rganic flux
Temperature
Salinity
Depth

D OS
GC M D OTS

0 0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8

Spearman Rank Correlation

B BioEnv results

%Sand
%Mud
Mean Grain size
% CaCO (Total)
Temperature
%G ravel
Salinity
% CaCO Mud
Depth
O rganic flux
GC M D OSM G 04-070-11

0 0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8

Spearman Rank Correlation

Figure 11. Spearman’s rank correlation for the relationship between benthic foraminiferal species abundances and environmental attributes
(A) and abundances of microbiota and the same environmental attributes (B). The correlation to individual parameters (striped bars) is
much lower than can be obtained with a multivariate approach (solid bars). The benthic foraminifera also show a much stronger correla-
tion to the environmental variables.

ly the same degree of variance (Ds = 0.799) as for all 6 variables,
indicating that these three factors are most strongly associated with
variations in the benthic assemblages across the sites. There is a
strong correlation between salinity and water depth (R2 = 0.80), and
the strong association between these two factors and the benthic
foraminiferal distributions emphasizes the influence of water depth
and associated variables on the foraminiferal species. The strength
of this relationship is consistent with studies in other settings, as
discussed later. The organic flux derived from the total number of
BF/g also correlates strongly with the species distributions. The
relationship between these three factors and the species distribution
is illustrated in Figure 12, with the physical variables overlain on
the species similarity plot.
Total Microbiota
Compared to the benthic foraminifera, the total microbiota have a
much lower correlation to the measured environmental variables
(Figure 11B). The highest correlation is Ds = 0.654, compared to Ds
= 0.802 for the benthic foraminifera. Interestingly, however, the
relationship to the environmental factors is similar for both analy-
ses. The microbiota are most strongly correlated to the factors %
gravel, %CaCO3, mud, depth, organic carbon flux, salinity and
mean grain size. These factors differ from the benthic foraminifera
only in the correlation to mean grain size, and the absence of a cor-
relation to temperature. This result implies that the response of the
benthic foraminiferal species to the environmental gradients is sim-

340
 BENTHIC FORAMS AS ENVIRONMENT INDICATORS

Figure 12. Similarity between sites based on the benthic foraminiferal assemblage (A), with the variation in key environmental parameters
across this distribution shown by the bubbles for salinity (B), water depth (C) and organic carbon flux (D). This multi-dimensional scale plot
is derived from the species similarity matrix, with the similarity in species composition between sites shown by their relative distance apart.
Salinity and water depth increase across the species distribution, with high salinity and greatest depths towards areas B and C. Organic car-
bon flux increases towards the lower left of the species distribution.

ilar to that of the whole micro-ecosystem. The stronger response of
the foraminiferal species to the environmental parameters, howev-
er, suggests that these organisms, when identified to species level,
may provide a more robust technique for benthic habitat mapping.
DISCUSSION
Taphonomic Preservation
environments. The most likely factors that affect the quantity of
degraded material in samples are in situ reworking, due to low sed-
imentation rates, or transport of material into (and out of) the sites.
Because the samples were collected using a Smith-McIntyre grab,
the foraminiferal population is likely to be somewhat biased
towards older material than a sample collected strictly from the sed-
iment surface. Abundant material of fresh appearance in some sam-
ples, however, indicates that the sampling method is not the sole
cause of the high relict quantity.

Overall Patterns In his discussion of living versus dead foraminiferal assem-

Evident from the study of preservation is the high proportion of
heavily modified and relict material present across the whole
region, but particularly abundant in Area A. For this study, it is
important to establish the likely cause of the high relict content, and
therefore whether the use of population data is valid in interpreting
blages, Murray (1982) suggests that a ratio of 1:1 would be expect-
ed in areas with normal sedimentation rates, while in areas of low
sedimentation, the proportion of living material would be far lower,
in the region of 1:10. An example quoted by Murray (1982) calcu-
lates rates of 1:4 to 1:6. In this study, although staining was not
undertaken as evidence of living populations, the fresh appearance

341
POST ET AL.
and light modification of specimens is used as an indication of the
relative proportion of in situ production and relict reworking.
The percentages of tests of fresh appearance plus light abra-
sion for both taxa in this study are <50% in all samples from Area
A, and typically below 20%, while in areas B and C, they are gen-
erally higher and often above 50%. These patterns suggest sub-
stantial differences in preservation between areas, possibly relat-
ed to differences in sedimentation rates. Dating of cores collected
in areas A and C support this pattern of higher sedimentation with-
in Area C compared to Area A. Vibrocores collected from Area A
indicate that Holocene carbonate material is present only in the
upper 1 m of most cores, with a transgressive peat sequence below
this (Harris et al., 2004). Vibrocore 21PC05 from Area A has a
lense of peat at 45 cm at the base of the Holocene carbonate
sequence. Radiocarbon dating of this peat deposit indicates a cor-
rected age of 9623 ± 71 years BP, and an age of 10,177 ± 49 years
BP recorded for peat deposited at 3.1 m in this core (Figure 13;
and the calibration based on Stuiver et al., 2005). The age of the
upper peat deposit indicates a sedimentation rate of 4.7 cm/1000
years within the carbonate sequence. Dating in Area C, based on
the foraminifera A. lessonii in gravity core 59GC17, yields a cor-
rected radiocarbon age of 475 ± 38 years BP at a depth of 10 cm,
with an age of 5655 ± 53 years BP at a depth of 2.95 m (Figure
13). The sedimentation rate for this core in Area C is 52 cm/1000
years, an order of magnitude higher than for the core in Area A.
These patterns support the interpretation that the foraminiferal
preservation types reflect the relative age of the surface sediments,
and are less affected by other chemical or physical processes
which may degrade the carbonate shells.
The wide range of preservation types in these sediments sug-
gests that the foraminifera represent a range of ages. This is prob-
ably the biggest constraint on comparison to modern physico-
chemical variables.
Differences between Areas
The presence of heavily reworked, large benthic foraminifera and
bivalve shells in the gravel fractions of samples from Area A (Harris
et al., 2004) suggests a high degree of reworking in situ, with long
residence times of the shells at, or near, the surface and little, or
slow, modern sedimentation (Murray, 1991). Cores from the region
also indicate low sedimentation rates (Harris et al., 2004, see above
discussion). Such features are not characteristic of sediments in
areas B and C. While bioerosion is present on lithoclasts from areas
B and C, the intensity is far lower and large, heavily-eroded bio-
clasts are absent from sediments.
The nature of the sediment also varies between the three areas.
The sediment in Area A has a much higher terrigenous content, par-
ticularly in the mud fraction (Harris et al., 2004). Sediments from
Area C typically have higher carbonate content. Lithics, where
present, are more commonly found in the sand fraction. Sediments
from Area B are transitional between areas A and C. Lithics in sed-
iment from Area C are found in samples from the channels (Harris
et al., 2004). These samples are also those which contain the most
abundant relict foraminifera, as would be expected in an environ-
ment which has been subject to high levels of sediment reworking
by currents.
342
The nature of the sedimentary environment can affect abrasion
in benthic organisms. A study of abrasion of gastropods and bivalves
in terrigenous sediments shows much more rapid abrasion than in
carbonate sediments due to the much more abrasive nature of silici-
clastic sediment (Driscoll and Weltin, 1973). Grain size also influ-
ences the effect, with the most rapid abrasion occurring in very
coarse sand, less in fine sand and least in medium sand. Mixed car-
bonate and siliciclastic sediments produced more rapid abrasion of
foraminifera than carbonate sediments (Moberly, 1968). Abrasion of
bivalves increases in either carbonate or mixed carbonate-siliciclas-
tic environments with hard grounds, compared to pure siliciclastic
environments (Best and Kidwell, 2000). The mixed siliciclastic-car-
bonates over Area A would therefore suggest intense abrasion of the
foraminiferal tests, while the dominantly fine-medium sand compo-
nent would lessen the impact of the abrasion. The high abrasion in
this area is likely to reflect a long residence time near the sediment
surface as a function of slower rates of accumulation.
Differences between Taxa
Experimental work to test the affects of abrasion on species preser-
vation indicates that susceptibility to damage varies between taxa.
After 2000 hours of tumbling (using foraminiferal specimens as the
abrasive), which is equivalent to transport of 34-100 km, 81% of A.
gibbosa tests were destroyed, compared to 68% of Q. tricarinata
tests (Peebles and Lewis, 1991). However, the Quinqueloculina
specimens were much more susceptible to dissolution than the
Amphistegina specimens. A number of the species of
Quinqueloculina included in this study are more fragile than
Amphistegina sp. due to their relatively smaller mass. Experimental
studies indicate that susceptibility to destruction by abrasion is con-
trolled by size (Peebles and Lewis, 1991). Examples of these more
fragile taxa include Q. subimpressa and Q. quinquecarinata.
As a result, the preservation index can be misleading because,
particularly in Amphistegina sp., fragmentation can result in a test
which rates as heavily degraded. In some cases, the surface preser-
vation may be excellent, suggesting that the fragmentation has
occurred relatively recently. This is quite different to a test which
has heavy surface abrasion. For example, the surface of the test
shown in Figure 2B is quite well-preserved although the degree of
breakage is very high. This is in contrast to the specimen in Figure
2D, which exhibits both surface degradation and test breakage. A
high level of breakage with relatively good surface preservation
was most evident in samples from Area C and may reflect the abun-
dance of sand-sized clastics in some samples. A future refinement
of the index, separating the effects of fragmentation from those of
surface degradation, would be useful.
Transport Pathways across Torres Strait
The foraminiferal dataset provides insight into the longer term sed-
iment transport across these areas, where the sediment samples in
Area A represent the history over several thousands of years. The
mechanical damage to foraminiferal tests collected across this study
region confirms the low sedimentation rates as indicated by dating.
Constraints provided by the species distributions suggest that most
of this reworking is occurring in situ, with very little cross-shelf
transport. Transportation pathways across this region are short and
are suggested by the angular surfaces on the sediment grains (Harris
 BENTHIC FORAMS AS ENVIRONMENT INDICATORS

Figure 13. Core logs for piston core 21PC05 (from Area A) and gravity core 59GC17 (from Area C). Radiocarbon dates are shown at the
collected depths, and are expressed as calibrated and reservoir corrected ages (based on Stuiver et al., 2005). The dates from piston core
21PC05 were derived from peat, while those from piston core 59GC17 are from benthic foraminifera.

343
POST ET AL.
et al., 2004). A previous study in the Fly River delta by Cole (1995)
indicated that foraminiferal transport occurs in bedload and sus-
pended sediment transport, with material carried by tidal currents
via the inter-reef channels. Evidence for this was shown through a
landward increase in planktonic foraminifera. Across the sample
sites, however, the relationship between planktonic foraminifera
and water depth is preserved, as it is also for many of the benthic
foraminiferal species.
The consistency of the relationship between water depth and
species abundance is particularly striking within areas B and C. The
shallower plateaux in these areas contain relatively higher abun-
dances of shallow water species such as A. ammoinoides and A.
papillosa (Figure 6A and B), while the adjacent channels contain
very low abundances of these species. The deeper water species,
such as S. pseudocarinata (Figure 6C), have relatively high abun-
dances in the channels and other deeper water sites, and extremely
low abundances across the plateaux. These patterns imply that the
faunas within areas B and C have been preserved in situ and mini-
mal transport within these areas has occurred. On this basis, the
relationship between the faunas and other environmental attributes
should be well defined, as corroborated by their consistency to pub-
lished relationships from other ocean basins (Murray, 1991; Van der
Zwann et al., 1999; Hayward et al., 2002) as discussed in the fol-
lowing section.
The low energy of the mid- to outer-shelf environments of
areas B and C as seen from the foraminiferal preservation patterns
and distributions is consistent with observations that suggest that
the currents across areas B and C, sourced from the upwelling of
Coral Sea water, are very clear with little sediment in suspension,
(Harris et al., 2004). The low sediment transport across Area C is
also shown by the muddy sand composition of sand waves sampled
within the area, suggesting that modern sand mobilization is not
occurring (Harris et al., 2004). The greater taphonomic decay of
tests within Area A is consistent with lower sedimentation rates in
the area, as shown also by radiocarbon dating.
Foraminiferal-Environment Relationships
Across the study sites, the benthic foraminiferal assemblages are
strongly correlated to changes in water depth, percent carbonate
mud, percent gravel, organic carbon flux, temperature and salinity.
These environmental preferences are consistent with those cited in
the literature for other ocean basins. In a study in the northern
Adriatic Sea, Jorissen et al. (1992) notes that the foraminiferal
species composition varies according to the downward flux of
organic matter and the associated oxygen levels, with areas which
experience seasonally high organic fluxes characterized by oppor-
tunistic species. Van der Zwann et al. (1999) note that species of
benthic foraminifera have a strong response to organic carbon flux,
depending on adequate oxygenation of the environment. The rela-
tionship between foraminifera and depth is considered to mostly
reflect the associated change in oxygen concentration (Van der
Zwann et al., 1999). However, other factors which may be depth
dependent, such as grain size, salinity and temperature can also be
important (Murray, 1991; Hayward et al., 2002). Grain size affects
the distribution of species in different environments due to the dif-
ferent habitat modes of foraminiferal species. Gravel and larger
344
sized components in the sediment, for instance, may provide a habi-
tat for those species which attach their tests to hard substrates, while
muddy sediments may allow a higher proportion of infaunal species
(Kitazato, 1994) The strong correlation between species distribu-
tions and factors such as grain size, organic flux, depth, temperature
and salinity in this study are therefore consistent with known rela-
tionships in other environments.
CONCLUSIONS
This study applied multivariate statistical techniques to determine
the relationships between benthic foraminifera and the whole
microbiotic community to changes in the environment. Both groups
exhibit a similar association to the environmental gradients.
However, the benthic foraminifera show a much stronger correla-
tion. The benthic foraminifera are strongly correlated to changes in
water depth, percent carbonate mud, percent gravel, organic carbon
flux, temperature and salinity. The application of a preservation
index, based on two foraminiferal genera, and taking into account
both fragmentation and dissolution, provides a good indication of
the degree of sediment reworking. However, the strength of the
foraminiferal-environment relationship, even at fine scales, sug-
gests that preservation is more strongly influenced by the low sedi-
mentation rates rather than sediment reworking through transport
across this margin. The consistency between species distributions
and environmental variables, despite the high degree of degradation
in many of the samples, provides much greater confidence in apply-
ing benthic habitat mapping techniques around the Australian mar-
gin, where sedimentation rates are low and reworking high.
ACKNOWLEDGMENTS
Our thanks to all those who provided the technical and scientific
support for the survey on RV Franklin. In particular, we thank Peter
Harris (Cruise leader, Geoscience Australia), Eric Madsen and
Bernie Heaney (CSIRO), Jon Stratton and Lyndon O’Grady
(Geoscience Australia) and Kevin Hooper (James Cook
University). Captain Ian Sinclair and the officers and crew of the
RV Franklin are also thanked for their assistance during the voyage.
Laboratory support was provided by Richard Brown, Tony Watson,
Alex McLachlan and Neil Ramsay (Sedimentology Laboratory,
Geoscience Australia). We thank Lynda Radke and Jon Clarke
(Geoscience Australia), Daniela Schmidt (University of Bristol) and
Robin Edwards (Trinity College, Dublin) for their constructive
reviews of the manuscript. This paper is published with the permis-
sion of the Executive Director, Geoscience Australia.
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APPENDIX I
Foraminiferal species list for abundance > 2 percent

Agglutinella arenata (Said) Nummulites venosus (Fitchel and Moll)

Ammomassilina alveoliniformis (Millet) Peneroplis pertusus (Forskål)
Ammonia spp. Plotnikovina timorea Loeblich and Tappan
Amphistegina papillosa Said Pseudogaudryina pacifica Cushman and McCulloch
Amphistegina spp. Pseudotriloculina patagonica (d'Orbigny)
Assilina ammonoides (Gronovius) Quinqueloculina crassicarinata Collins
Asterorotalia gaimardi (d'Orbigny) Quinqueloculina incisa Vella
Cibicides refulgens (Montfort) Quinqueloculina philippinensis Cushman
Cibicides type Quinqueloculina spp.
Cibicidoides basilanensis McCulloch Reussella spp.
Discorbia type Rosalina spp.
Elphidium crispum (Linné) Siphoniferoides siphoniferous (Brady)
Elphidium spp. Siphotextularia mestayerae Vella
Gaudryina spp. Spiroloculina subimpressa Parr
Hanzawaia spp. Spiroplectinella pseudocarinata (Cushman)
Heterolepa subhaidingeri (Parr) Spirosigmoilina parri Collins
Loxostima spp. Textularia cushmani Said
Melonis spp. Textularia foliacea Heron-Allen and Earland
Miniacina miniacea (Pallas) Textularia secasensis Lablicker and McCulloch
Neoconorbina spp. Textularia stricta Cushman
Neoeponides bradyi LeCalvez Textularia truncata Höglund
Neouvigerina spp. Textularia spp.
Nubeculina advena Cushman Triloculina pseudooblonga (Zheng)

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APPENDIX II
Preservation levels of Amphistegina spp.

Sample % % % % % Tot. no. Sample % % % % % Tot. no.

no. fresh light moderate heavy relict analyzed no. fresh light moderate heavy relict analyzed

Area A Area C
234/1GR1A 1.92 11.54 28.85 57.69 0.00 52 234/54GR43A 50.00 0.00 0.00 50.00 0.00 2
234/2GR2A 3.95 14.47 32.89 47.37 1.32 76 234/55GR44A 16.67 20.83 37.50 25.00 0.00 24
234/3GR3A 2.11 6.32 23.16 48.42 20.00 95 234/57GR45A 20.95 21.90 22.86 27.62 6.67 105
234/4GR4A 5.71 5.71 14.29 65.71 8.57 35 234/58GR47A 0.00 25.00 25.00 25.00 25.00 4
234/5GR5A 2.33 11.63 27.91 56.98 1.16 86 234/59GR48A 19.57 36.96 19.57 23.91 0.00 46
234/6GR6A 6.02 8.43 25.30 56.63 3.61 83 234/60GR49A 0.00 66.67 0.00 0.00 33.33 3
234/7GR7A 3.28 6.56 21.31 68.85 0.00 61 234/61GR50A 0.00 20.00 0.00 60.00 20.00 5
234/8GR8A 2.35 16.47 34.12 41.18 5.88 85 234/62GR51A 11.11 11.11 66.67 11.11 0.00 9
234/9GR9A 6.03 12.07 30.17 51.72 0.00 116 234/63GR52A 0.00 25.00 0.00 75.00 0.00 4
234/10GR10A 4.00 8.00 26.67 56.00 5.33 75 234/64GR53A 6.90 3.45 13.79 44.83 31.03 29
234/11GR11A 4.17 13.89 27.78 54.17 0.00 72 234/65GR54A 56.86 19.61 15.69 7.84 0.00 51
234/12GR12A 0.00 2.83 21.70 43.40 32.08 106 234/66GR55A 14.63 4.88 29.27 46.34 4.88 41
234/13GR13A 3.57 8.93 25.00 58.93 3.57 56 234/67GR56A 6.67 20.00 40.00 33.33 0.00 15
234/14GR14A 3.51 14.04 17.54 42.11 22.81 57 234/68GR57A 13.51 18.92 16.22 48.65 2.70 37
234/15GR15A 2.75 13.76 31.19 52.29 0.00 109 234/69GR58A 9.09 0.00 9.09 45.45 36.36 11
234/16GR16A 3.39 3.39 20.34 69.49 3.39 59 234/70GR59A 11.76 31.37 19.61 25.49 11.76 51
234/17GR17A 8.96 19.40 31.34 34.33 5.97 67 234/72GR60A 16.67 8.33 25.00 33.33 16.67 12
234/18GR18A 0.00 1.22 24.39 45.12 29.27 82 234/73GR61A 0.00 40.00 40.00 20.00 0.00 5
234/19GR19A 10.14 13.04 27.54 49.28 0.00 69 234/74GR62A 23.81 9.52 9.52 47.62 9.52 21
234/20GR20A 0.00 2.70 13.51 55.41 28.38 74 234/75GR63A 8.33 0.00 25.00 58.33 8.33 12
234/21GR21A 2.78 23.61 34.72 37.50 1.39 72

Area B
234/28GR22A 0.00 12.90 25.81 61.29 0.00 31
234/29GR23A 15.52 29.31 36.21 17.24 1.72 58
234/30GR24A 23.61 23.61 25.00 26.39 1.39 72
234/31GR25A 14.06 10.94 37.50 34.38 3.13 64
234/32GR26A 0.00 14.29 28.57 57.14 0.00 7
234/33GR27A 16.48 23.08 19.78 37.36 3.30 91
234/34GR28A 8.57 2.86 11.43 57.14 20.00 35
234/35GR29A 0.00 12.00 24.00 48.00 16.00 25
234/36GR30A 4.35 8.70 21.74 65.22 0.00 23
234/37GR31A 10.45 14.18 14.93 39.55 20.90 134
234/38GR32A 0.00 66.67 0.00 33.33 0.00 3
234/39GR33A 11.90 11.90 30.95 40.48 4.76 42
234/40GR34A 7.14 11.43 21.43 55.71 4.29 70
234/41GR35A 22.22 22.22 22.22 33.33 0.00 9
234/42GR36A 0.00 15.63 21.88 56.25 6.25 32
234/43GR37A 7.89 13.16 21.05 50.00 7.89 38
234/44GR38A 4.55 0.00 9.09 50.00 36.36 22
234/45GR39A 0.00 20.00 40.00 40.00 0.00 10
234/46GR40A 10.00 13.33 40.00 36.67 0.00 30
234/47GR41A 15.38 8.97 37.18 37.18 1.28 78
234/48GR42A 10.08 10.08 24.37 50.42 5.04 119

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APPENDIX III
Preservation levels of Quinqueloculina spp.

Sample % % % % % Tot. no. Sample % % % % % Tot. no.

no. fresh light moderate heavy relict analyzed no. fresh light moderate heavy relict analyzed

Area A Area C
234/1GR1A 0.00 13.64 13.64 50.00 22.73 22 234/54GR43A 0.00 11.76 11.76 29.41 47.06 17
234/2GR2A 1.69 6.78 16.95 57.63 16.95 59 234/55GR44A 9.09 9.09 45.45 36.36 0.00 11
234/3GR3A 0.00 0.00 14.71 58.82 26.47 34 234/57GR45A 6.67 6.67 20.00 33.33 33.33 15
234/4GR4A 3.13 3.13 9.38 78.13 6.25 32 234/58GR46A 0.00 15.38 15.38 69.23 0.00 13
234/5GR5A 1.27 15.19 26.58 46.84 10.13 79 234/58GR47A 9.09 27.27 27.27 36.36 0.00 11
234/6GR6A 1.75 14.04 17.54 54.39 12.28 57 234/59GR48A 21.43 7.14 14.29 28.57 28.57 14
234/7GR7A 0.00 28.00 12.00 60.00 0.00 25 234/60GR49A 15.38 11.54 26.92 23.08 23.08 26
234/8GR8A 2.78 5.56 22.22 63.89 5.56 36 234/61GR50A 25.81 12.90 22.58 19.35 19.35 31
234/9GR9A 0.00 0.00 23.08 61.54 15.38 26 234/62GR51A 5.36 12.50 28.57 42.86 10.71 56
234/10GR10A 2.22 8.89 15.56 57.78 15.56 45 234/63GR52A 8.33 25.00 0.00 33.33 33.33 12
234/11GR11A 4.76 14.29 33.33 47.62 0.00 21 234/64GR53A 4.00 12.00 4.00 20.00 60.00 25
234/12GR12A 0.00 3.70 3.70 48.15 44.44 27 234/65GR54A 24.00 28.00 32.00 16.00 0.00 25
234/13GR13A 0.00 6.45 16.13 41.94 35.48 31 234/66GR55A 2.94 2.94 23.53 29.41 41.18 34
234/14GR14A 2.13 6.38 21.28 55.32 14.89 47 234/67GR56A 0.00 9.09 18.18 63.64 9.09 22
234/15GR15A 2.50 12.50 27.50 50.00 7.50 40 234/68GR57A 12.77 23.40 21.28 29.79 12.77 47
234/16GR16A 0.00 4.55 18.18 56.82 20.45 44 234/69GR58A 11.76 23.53 5.88 23.53 35.29 17
234/17GR17A 3.13 0.00 31.25 59.38 6.25 32 234/70GR59A 0.00 0.00 25.00 25.00 50.00 4
234/18GR18A 0.00 8.89 35.56 40.00 15.56 45 234/72GR60A 10.53 0.00 10.53 36.84 42.11 19
234/19GR19A 4.35 10.87 17.39 65.22 2.17 46 234/73GR61A 12.12 6.06 30.30 30.30 21.21 33
234/20GR20A 3.45 3.45 44.83 48.28 0.00 29 234/74GR62A 0.00 0.00 4.55 36.36 59.09 22
234/21GR21A 0.00 6.06 18.18 66.67 9.09 33 234/75GR63A 0.00 13.64 0.00 45.45 40.91 22

Area B
234/28GR22A 8.51 14.89 23.40 48.94 4.26 47
234/29GR23A 6.67 46.67 0.00 46.67 0.00 15
234/30GR24A 13.64 18.18 22.73 45.45 0.00 22
234/31GR25A 0.00 11.76 11.76 47.06 29.41 17
234/32GR26A 9.52 33.33 14.29 38.10 4.76 21
234/33GR27A 18.75 6.25 12.50 50.00 12.50 16
234/34GR28A 2.70 0.00 13.51 35.14 48.65 37
234/35GR29A 0.00 10.71 21.43 50.00 17.86 28
234/36GR30A 0.00 9.09 22.73 63.64 4.55 22
234/37GR31A 10.53 10.53 26.32 36.84 15.79 19
234/38GR32A 0.00 0.00 37.50 25.00 37.50 8
234/39GR33A 4.55 13.64 9.09 27.27 45.45 22
234/40GR34A 13.04 13.04 21.74 52.17 0.00 23
234/41GR35A 0.00 33.33 0.00 50.00 16.67 6
234/42GR36A 12.90 3.23 16.13 45.16 22.58 31
234/43GR37A 6.90 13.79 31.03 34.48 13.79 29
234/44GR38A 5.56 16.67 33.33 38.89 5.56 18
234/45GR39A 6.82 20.45 22.73 43.18 6.82 44
234/46GR40A 10.00 20.00 10.00 60.00 0.00 10
234/47GR41A 0.00 9.09 18.18 54.55 18.18 11
234/48GR42A 0.00 6.25 18.75 43.75 31.25 32

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