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

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BENTHIC FORAMS AS ENVIRONMENT INDICATORS

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 Survey 234 aboard RV Franklin in January-February 2002 (Figure
were generally flat and included some rocky and reef environments. 1). The physical characteristics of each site were determined
through several analyses. Surface grab samples were analyzed for
Area C is located 105 to 120 km offshore and comprises a sub- gravel, sand and silt content by washing through 2 mm and 63 μm
marine valley system aligned west-northwest–east-northeast sieves, and the <2 mm fraction was analyzed by laser particle size.
(Figure 1). The valley occurs as two separate limbs with the deeper Carbonate content was determined on the mud and sand fractions
southern channels representing relict features formed by tidal cur- using the carbonate bomb method. In addition, the seabed morphol-
rent scour during Pleistocene sea level lowstands (Heap et al., ogy was characterized based on swath bathymetry data, together
2003). The water depth at sampling sites across this area ranges with seabed photography. Temperature and salinity measurements
from 50 to 220 m, and the shallow areas are mapped as platform were obtained at each site via conductivity/temperature/depth
reefs. The channels across both areas B and C act as conduits for (CTD) casts. The measurements from nearest the seabed were used
upwelled Coral Sea water, which is relatively cool, saline and nutri- in comparison with the benthic foraminiferal assemblages. Details
ent-rich (Harris et al., 2004). of each of these analyses are available in Harris et al. (2004).

Bottom measurements of temperature and salinity at each site Analysis of Benthic Foraminifera
indicate that each of these parameters is tightly correlated with
changes in water depth. As water depth increases, the temperature of Surface sediments were collected using a Smith-McIntyre grab. The
the bottom water decreases logarithmically (R2 = 0.75) while salini- grab material was dry-sieved to the 150 :m to 2 mm size fraction
ty increases (R2 = 0.80). The bottom water temperature ranges and split into aliquots, each containing at least 300 specimens of
between 25 and 29°C across all sites, and salinity between 34.33 and benthic foraminifera. The foraminifera were identified using a
35.18‰. The increase in salinity with depth is associated with binocular light microscope and referencing the taxonomic studies of
upwelling of saline, cool water from the Coral Sea into deeper chan- Hottinger et al. (1993), Loeblich and Tappan (1994), Hayward et al.
nels of areas B and C (Harris et al., 2004), while shallower nearshore (1997) and Haig (2002). The most abundant forms (>2% abundance)
areas are influenced by fresh water input from the Fly River. were identified to species level (see species list in Appendix I), while
rarer forms were grouped and identified to genera.
METHODS
Taphonomy
Analysis of Environmental Parameters
Two foraminiferal genera were selected for assessment of taphono-
Surface sediment samples were collected from 64 sites within the my, following an overview of preservation in samples from all three
Torres Strait–Gulf of Papua region during Geoscience Australia areas. Both genera selected: Amphistegina sp. and Quinqueloculina

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POST ET AL.

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

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
Figure 4. Site similarity (MDS) based on the abundance of micro- 55 m), S. pseudocarinata (maximum at 110 m) and H. subhaidin-
biota (measured as number of specimens/gram of sediment). geri (maximum at 85 m). These species tend to increase in abun-
Similarity is plotted as the relative distance between sites. Sites dance across Area B and shallower parts of Area C.
within Area A show a distinct difference in microbiota abundances Pseudogaudryina pacifica maintains its presence across the deep-
to areas B and C. There is substantial overlap in site similarities er water sites and has an abundance of 6%. Cibicides-type species
within areas B and C. and C. refulgens are distributed across all water depths and the
highest abundance occurs at depths >120 m for Cibicides-type
potential for mapping the ecosystem through detailed studies of species. The maximum abundance of C. refulgens occurs at 90 m.
species within this phylum. However, it maintains high abundances across depths >120 m.
These two species are most common within Area C.
Foraminiferal Abundances
The distribution of several of these key species across the sites
Benthic foraminifera are abundant in each of the grab samples is shown in Figures 6A-C, illustrating the relationship between
examined. Thirty-one species >2% and minor species (numerical) bathymetry and species abundances on a site-by-site basis. Assilina

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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.

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.

ammonoides (Figure 6A) has consistently high abundances across Taphonomic Features of Benthic Foraminifera
Area A, while high abundances of this species within areas B and C
are restricted to the shallower plateaux, and very low abundances The taphonomic changes in selected genera of benthic foraminifera
occur at deeper sites. A similar distribution is shown for A. papil- have been analyzed across all the sites. All samples show evidence
losa in Figure 6B, with the highest abundance of this species in of some form of mechanical or chemical breakdown, but with vari-
areas B and C occurring at the shallowest sites. Spiroplectinella able intensity between sites. Effects observed include pitting, break-
pseudocarinata (Figure 6C), by contrast, shows highest abundances age, scalloping of test edges, flattening or removal of surface orna-
at the deepest sites, and therefore has very low abundance at sites in mentation or surface layers, polishing, test infill and staining
areas A and B, with slightly increased numbers occurring at the few (Figure 2). Abrasion features are the most common form of test
relatively deeper sites in Area B. alteration, indicating mechanical breakdown due to bombardment
with fine-grained material. Dissolution effects, mainly seen as
Cluster analysis of the foraminiferal species data reveals 7 opacity of the test surface, are also common. The very heavy abra-
main clusters. Cluster 1 consists of just one sample from Area A, sion of some specimens indicates their long residence time near the
Site 4, with a similarity to other sites of 50%. Other sites from Area surface. There appears to be no correlation between species abun-
A form Cluster 2 (Figure 7), with similarities ranging between 70 dances and their preservation, implying that preservation is not a
and 90%. The foraminiferal assemblage at Site 4 contains higher function of species production or transportation to the sites.
abundances of Quinqueloculina philippinensis and Cibicides-type
species, and much lower abundances of N. venosus compared to the The preservation of Amphistegina sp. is poor in Area A, sug-
assemblages at other sites within Area A. Clusters in areas B and C gesting a high degree of reworking in these sediments. Areas B and
show some overlap between the two areas, with Cluster 4 the only C, however, are dominated by sites with >25% of tests with a fresh
distinct cluster, composed of sites from the deeper parts of Area C appearance or lightly modified (Figure 8). Across the remaining
(Figure 7). Similarities between sites within these remaining clus- sites, most of the tests fall into the heavily modified or relict cate-
ters range between 60-85%. gories. Average preservation values of Amphistegina sp. across all

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BENTHIC FORAMS AS ENVIRONMENT INDICATORS

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 all sites are shown in Figure 10. Details of Quinqueloculina sp.
preservation for Amphistegina sp. is provided in Appendix II. preservation are provided in Appendix III. In Area A, samples are
dominated by material that has been heavily degraded, >40-78%.
In Area A, samples are generally dominated by tests showing Relict content is low (0-20%) to moderate (>20-45%). The content
heavy abrasion, typically >40-70%. The abundance of both relict of material of fresh appearance is low throughout, <5%, with 12 of
and material of fresh appearance is generally low, <10%; 6 samples the 21 samples having no specimens. The proportion of lightly
have no relict specimens, while 5 of the 21 samples show values modified material is low, generally in the range 0-20%, with 9 sam-
>20%. Similarly, the abundance of material of fresh appearance is ples having no lightly modified specimens.
low, 0-10%; 3 samples have no tests of fresh appearance.
Area B samples are generally dominated by material with
Area B shows greater variability. Samples are generally domi- heavy degradation, 30-50%, although some samples show higher
nated by tests that are heavily degraded, typically in the range >25- levels of either relict or moderately abraded material. There is
65%. The relict content is variable; 7 samples have no relict speci- greater variability in the relict content where 14 out of the 21 sam-
mens, the remainder range from 1-35%. However, the percentage of ples have low values (0-20%) and the remainder range from 20 to
material of fresh appearance and light modification is generally 49%. The proportion of material of fresh appearance is similarly
higher than in Area A. Area C also shows a high level of variabili- variable, 0-18%; 14 of 21 samples have no specimens with a fresh
ty. About half of the samples are dominated by tests in the heavily appearance. Where present, the proportion of material of fresh
modified category, although the dominance is much less clear cut appearance is generally higher than in Area A samples, as is the pro-
than in samples from areas A and B. The remaining samples are portion of lightly modified material.
dominated by light or moderate abrasion, or, in one case, by mate-
rial of fresh appearance. In Area C, samples show greater variability; some contain a
higher content (up to 50%) of material that has a fresh appearance
The preservation of Quinqueloculina sp. is poor throughout; and light modification, with little or no relict material; others are
sites with poor preservation outnumber sites where well-preserved biased towards being heavily abraded and relict, and contain lit-
material dominates (Figure 8). Average preservation values across tle material with fresh appearance or light modification. Relict

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POST ET AL.

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.

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

Several key environmental attributes vary between the species clus- Analysis of individual parameters indicates that depth has a strong
ters (Table 4). Mean water depth varies substantially between the 7 correlation to the site similarity expressed by the benthic assem-
clusters, ranging from 24.5 m in Cluster 1 to 119 m in Cluster 4 blages, with a Spearman’s rank correlation coefficient (Ds) of 0.68
(Table 4). The grain size and carbonate content also show a large (Figure 11A). Stronger correlations were found when factors were
range in values across the sites. The number of benthic combined, with the best correlation observed for the combination of
foraminifera/g (BF/g) indicates a high organic flux in Cluster 2 % gravel, % CaCO3 mud, depth, organic carbon flux, temperature
(Area A), compared to the exceptionally low flux in Cluster 1 (Site and salinity (Ds = 0.802). However, the combination of just three
4, also Area A). The organic flux is more moderate across the 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, Total Microbiota
indicating that these three factors are most strongly associated with
variations in the benthic assemblages across the sites. There is a Compared to the benthic foraminifera, the total microbiota have a
strong correlation between salinity and water depth (R2 = 0.80), and much lower correlation to the measured environmental variables
the strong association between these two factors and the benthic (Figure 11B). The highest correlation is Ds = 0.654, compared to Ds
foraminiferal distributions emphasizes the influence of water depth = 0.802 for the benthic foraminifera. Interestingly, however, the
and associated variables on the foraminiferal species. The strength relationship to the environmental factors is similar for both analy-
of this relationship is consistent with studies in other settings, as ses. The microbiota are most strongly correlated to the factors %
discussed later. The organic flux derived from the total number of gravel, %CaCO3, mud, depth, organic carbon flux, salinity and
BF/g also correlates strongly with the species distributions. The mean grain size. These factors differ from the benthic foraminifera
relationship between these three factors and the species distribution only in the correlation to mean grain size, and the absence of a cor-
is illustrated in Figure 12, with the physical variables overlain on relation to temperature. This result implies that the response of the
the species similarity plot. 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 environments. The most likely factors that affect the quantity of
the foraminiferal species to the environmental parameters, howev- degraded material in samples are in situ reworking, due to low sed-
er, suggests that these organisms, when identified to species level, imentation rates, or transport of material into (and out of) the sites.
may provide a more robust technique for benthic habitat mapping. Because the samples were collected using a Smith-McIntyre grab,
the foraminiferal population is likely to be somewhat biased
DISCUSSION 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
Taphonomic Preservation
cause of the high relict quantity.

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


blages, Murray (1982) suggests that a ratio of 1:1 would be expect-
Evident from the study of preservation is the high proportion of ed in areas with normal sedimentation rates, while in areas of low
heavily modified and relict material present across the whole sedimentation, the proportion of living material would be far lower,
region, but particularly abundant in Area A. For this study, it is in the region of 1:10. An example quoted by Murray (1982) calcu-
important to establish the likely cause of the high relict content, and lates rates of 1:4 to 1:6. In this study, although staining was not
therefore whether the use of population data is valid in interpreting 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 The nature of the sedimentary environment can affect abrasion
relative proportion of in situ production and relict reworking. in benthic organisms. A study of abrasion of gastropods and bivalves
in terrigenous sediments shows much more rapid abrasion than in
The percentages of tests of fresh appearance plus light abra- carbonate sediments due to the much more abrasive nature of silici-
sion for both taxa in this study are <50% in all samples from Area clastic sediment (Driscoll and Weltin, 1973). Grain size also influ-
A, and typically below 20%, while in areas B and C, they are gen- ences the effect, with the most rapid abrasion occurring in very
erally higher and often above 50%. These patterns suggest sub- coarse sand, less in fine sand and least in medium sand. Mixed car-
stantial differences in preservation between areas, possibly relat- bonate and siliciclastic sediments produced more rapid abrasion of
ed to differences in sedimentation rates. Dating of cores collected foraminifera than carbonate sediments (Moberly, 1968). Abrasion of
in areas A and C support this pattern of higher sedimentation with- bivalves increases in either carbonate or mixed carbonate-siliciclas-
in Area C compared to Area A. Vibrocores collected from Area A tic environments with hard grounds, compared to pure siliciclastic
indicate that Holocene carbonate material is present only in the environments (Best and Kidwell, 2000). The mixed siliciclastic-car-
upper 1 m of most cores, with a transgressive peat sequence below bonates over Area A would therefore suggest intense abrasion of the
this (Harris et al., 2004). Vibrocore 21PC05 from Area A has a foraminiferal tests, while the dominantly fine-medium sand compo-
lense of peat at 45 cm at the base of the Holocene carbonate nent would lessen the impact of the abrasion. The high abrasion in
sequence. Radiocarbon dating of this peat deposit indicates a cor- this area is likely to reflect a long residence time near the sediment
rected age of 9623 ± 71 years BP, and an age of 10,177 ± 49 years surface as a function of slower rates of accumulation.
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 Differences between Taxa
upper peat deposit indicates a sedimentation rate of 4.7 cm/1000
years within the carbonate sequence. Dating in Area C, based on Experimental work to test the affects of abrasion on species preser-
the foraminifera A. lessonii in gravity core 59GC17, yields a cor- vation indicates that susceptibility to damage varies between taxa.
rected radiocarbon age of 475 ± 38 years BP at a depth of 10 cm, After 2000 hours of tumbling (using foraminiferal specimens as the
with an age of 5655 ± 53 years BP at a depth of 2.95 m (Figure abrasive), which is equivalent to transport of 34-100 km, 81% of A.
13). The sedimentation rate for this core in Area C is 52 cm/1000 gibbosa tests were destroyed, compared to 68% of Q. tricarinata
years, an order of magnitude higher than for the core in Area A. tests (Peebles and Lewis, 1991). However, the Quinqueloculina
These patterns support the interpretation that the foraminiferal specimens were much more susceptible to dissolution than the
preservation types reflect the relative age of the surface sediments, Amphistegina specimens. A number of the species of
and are less affected by other chemical or physical processes Quinqueloculina included in this study are more fragile than
which may degrade the carbonate shells. Amphistegina sp. due to their relatively smaller mass. Experimental
studies indicate that susceptibility to destruction by abrasion is con-
The wide range of preservation types in these sediments sug- trolled by size (Peebles and Lewis, 1991). Examples of these more
gests that the foraminifera represent a range of ages. This is prob- fragile taxa include Q. subimpressa and Q. quinquecarinata.
ably the biggest constraint on comparison to modern physico-
chemical variables. As a result, the preservation index can be misleading because,
particularly in Amphistegina sp., fragmentation can result in a test
Differences between Areas which rates as heavily degraded. In some cases, the surface preser-
vation may be excellent, suggesting that the fragmentation has
The presence of heavily reworked, large benthic foraminifera and occurred relatively recently. This is quite different to a test which
bivalve shells in the gravel fractions of samples from Area A (Harris has heavy surface abrasion. For example, the surface of the test
et al., 2004) suggests a high degree of reworking in situ, with long shown in Figure 2B is quite well-preserved although the degree of
residence times of the shells at, or near, the surface and little, or breakage is very high. This is in contrast to the specimen in Figure
slow, modern sedimentation (Murray, 1991). Cores from the region 2D, which exhibits both surface degradation and test breakage. A
also indicate low sedimentation rates (Harris et al., 2004, see above high level of breakage with relatively good surface preservation
discussion). Such features are not characteristic of sediments in was most evident in samples from Area C and may reflect the abun-
areas B and C. While bioerosion is present on lithoclasts from areas dance of sand-sized clastics in some samples. A future refinement
B and C, the intensity is far lower and large, heavily-eroded bio- of the index, separating the effects of fragmentation from those of
clasts are absent from sediments. surface degradation, would be useful.

The nature of the sediment also varies between the three areas. Transport Pathways across Torres Strait
The sediment in Area A has a much higher terrigenous content, par-
ticularly in the mud fraction (Harris et al., 2004). Sediments from The foraminiferal dataset provides insight into the longer term sed-
Area C typically have higher carbonate content. Lithics, where iment transport across these areas, where the sediment samples in
present, are more commonly found in the sand fraction. Sediments Area A represent the history over several thousands of years. The
from Area B are transitional between areas A and C. Lithics in sed- mechanical damage to foraminiferal tests collected across this study
iment from Area C are found in samples from the channels (Harris region confirms the low sedimentation rates as indicated by dating.
et al., 2004). These samples are also those which contain the most Constraints provided by the species distributions suggest that most
abundant relict foraminifera, as would be expected in an environ- of this reworking is occurring in situ, with very little cross-shelf
ment which has been subject to high levels of sediment reworking transport. Transportation pathways across this region are short and
by currents. are suggested by the angular surfaces on the sediment grains (Harris

342
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) sized components in the sediment, for instance, may provide a habi-
indicated that foraminiferal transport occurs in bedload and sus- tat for those species which attach their tests to hard substrates, while
pended sediment transport, with material carried by tidal currents muddy sediments may allow a higher proportion of infaunal species
via the inter-reef channels. Evidence for this was shown through a (Kitazato, 1994) The strong correlation between species distribu-
landward increase in planktonic foraminifera. Across the sample tions and factors such as grain size, organic flux, depth, temperature
sites, however, the relationship between planktonic foraminifera and salinity in this study are therefore consistent with known rela-
and water depth is preserved, as it is also for many of the benthic tionships in other environments.
foraminiferal species.
CONCLUSIONS
The consistency of the relationship between water depth and
species abundance is particularly striking within areas B and C. The This study applied multivariate statistical techniques to determine
shallower plateaux in these areas contain relatively higher abun- the relationships between benthic foraminifera and the whole
dances of shallow water species such as A. ammoinoides and A. microbiotic community to changes in the environment. Both groups
papillosa (Figure 6A and B), while the adjacent channels contain exhibit a similar association to the environmental gradients.
very low abundances of these species. The deeper water species, However, the benthic foraminifera show a much stronger correla-
such as S. pseudocarinata (Figure 6C), have relatively high abun- tion. The benthic foraminifera are strongly correlated to changes in
dances in the channels and other deeper water sites, and extremely water depth, percent carbonate mud, percent gravel, organic carbon
low abundances across the plateaux. These patterns imply that the flux, temperature and salinity. The application of a preservation
faunas within areas B and C have been preserved in situ and mini- index, based on two foraminiferal genera, and taking into account
mal transport within these areas has occurred. On this basis, the both fragmentation and dissolution, provides a good indication of
relationship between the faunas and other environmental attributes the degree of sediment reworking. However, the strength of the
should be well defined, as corroborated by their consistency to pub- foraminiferal-environment relationship, even at fine scales, sug-
lished relationships from other ocean basins (Murray, 1991; Van der gests that preservation is more strongly influenced by the low sedi-
Zwann et al., 1999; Hayward et al., 2002) as discussed in the fol- mentation rates rather than sediment reworking through transport
lowing section. across this margin. The consistency between species distributions
and environmental variables, despite the high degree of degradation
The low energy of the mid- to outer-shelf environments of in many of the samples, provides much greater confidence in apply-
areas B and C as seen from the foraminiferal preservation patterns ing benthic habitat mapping techniques around the Australian mar-
and distributions is consistent with observations that suggest that gin, where sedimentation rates are low and reworking high.
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
ACKNOWLEDGMENTS
also shown by the muddy sand composition of sand waves sampled Our thanks to all those who provided the technical and scientific
within the area, suggesting that modern sand mobilization is not support for the survey on RV Franklin. In particular, we thank Peter
occurring (Harris et al., 2004). The greater taphonomic decay of Harris (Cruise leader, Geoscience Australia), Eric Madsen and
tests within Area A is consistent with lower sedimentation rates in Bernie Heaney (CSIRO), Jon Stratton and Lyndon O’Grady
the area, as shown also by radiocarbon dating. (Geoscience Australia) and Kevin Hooper (James Cook
University). Captain Ian Sinclair and the officers and crew of the
Foraminiferal-Environment Relationships RV Franklin are also thanked for their assistance during the voyage.
Laboratory support was provided by Richard Brown, Tony Watson,
Across the study sites, the benthic foraminiferal assemblages are Alex McLachlan and Neil Ramsay (Sedimentology Laboratory,
strongly correlated to changes in water depth, percent carbonate Geoscience Australia). We thank Lynda Radke and Jon Clarke
mud, percent gravel, organic carbon flux, temperature and salinity. (Geoscience Australia), Daniela Schmidt (University of Bristol) and
These environmental preferences are consistent with those cited in Robin Edwards (Trinity College, Dublin) for their constructive
the literature for other ocean basins. In a study in the northern reviews of the manuscript. This paper is published with the permis-
Adriatic Sea, Jorissen et al. (1992) notes that the foraminiferal sion of the Executive Director, Geoscience Australia.
species composition varies according to the downward flux of
organic matter and the associated oxygen levels, with areas which REFERENCES
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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)

345
POST ET AL.

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

346
BENTHIC FORAMS AS ENVIRONMENT INDICATORS

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

347
POST ET AL.

348