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Estuarine, Coastal and Shelf Science 58 (2003) 677–697

The origin of modern agglutinated foraminiferalassemblages: evidence from a stratified fjord

John W. Murraya,*, Elisabeth Alveb, Andrew Cundyc

aSchool of Ocean and Earth Science, Southampton Oceanography Centre, European Way, Southampton SO14 3ZH, UKbDepartment of Geology, University of Oslo, PO Box 1047 Blindern, N0316 Oslo, Norway

cSchool of Chemistry, Physics and Environmental Science, University of Sussex, Falmer, Brighton, BN1 9QJ, UK

Received 17 March 2003; accepted 4 June 2003

Abstract

Loch Etive, a silled 145 m deep fjord on the Scottish west coast, provides an example of modern benthic foraminiferal

assemblages at intermediate depths (i.e., below the intertidal zone and above the CCD) consisting almost exclusively of organic-cemented agglutinated forms. Since such faunas from intermediate depths are rare in modern oceans but relatively common in thefossil record, the present study allows new insights into one kind of ancient environment for which there are few modern analogues.

The strong dominance of agglutinated forms (both living and in some dead assemblages of foraminifera to the exclusion ofcalcareous taxa) is attributed to the unusual oceanographic conditions. These include a combination of restricted deep-waterrenewals and strong influence of freshwater which drains through large areas (relative to the size of the loch) of vegetated land. The

result is calm bottom water conditions with commonly occurring oxygen depletion (although not anoxic), brackish waterthroughout the water column (salinity 28 in the deeper parts), and organic-rich (mostly terrestrially derived) sediments withgeochemical properties, which, to a much larger degree than open marine ones, are controlled by local input. This environmentsupports low abundance and low diversity live assemblages, mainly restricted to the surface 1 cm of sediment. The dead assemblages

show similar faunal characteristics, but the calcareous components are, due to carbonate dissolution, even more reduced. One of thecalcareous species in Loch Etive is Elphidium albiumbilicatum. Its occurrence is the first record in British waters and it matches thepreviously suggested southern limit of its distribution.

Analysis of a 90 cm long core representing sediments deposited over the past two centuries shows the presence of a calcareousdominated assemblage, including more marine species, with a higher diversity, in the lower part. This suggests that Loch Etive isin the process of going from a marine, to a more terrestrial dominated environment. The relatively high sedimentation rate (0.5 cm

per yr), the apparent lack of smearing through bioturbation, and the presence of faunal changes in response to reduced marineinfluence over the past centuries, shows that Loch Etive has a good potential for performing high-resolution climatic studies.� 2003 Elsevier Ltd. All rights reserved.

Keywords: agglutinated foraminifera; Loch Etive; fjord; oxygen depletion; carbonate dissolution; climatic history

1. Introduction

In the fossil record there are numerous records ofexclusively agglutinated assemblages of foraminiferathat are inferred to have lived at intermediate waterdepths. In modern environments such exclusively

* Corresponding author.

E-mail addresses: [email protected] (J.W. Murray),

[email protected] (E. Alve), [email protected] (A. Cundy).

0272-7714/03/$ - see front matter � 2003 Elsevier Ltd. All rights reserved.

doi:10.1016/S0272-7714(03)00179-3

agglutinated (living) assemblages occur to a limitedextent in parts of brackish estuaries, fjords, and shallowbays or lagoons but they are mainly confined tointertidal salt marshes and to open oceanic areas belowthe CCD most of which are abyssal (Schroder et al.,1988) or several hundred metres deep (as around Antarc-tica, e.g., Ward et al., 1986). Yet it is clear that the fossilexamples do not always represent these extremes. It istherefore of considerable interest to discover furthermodern environments in which agglutinated forms makeup a very high proportion of the assemblages and which

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678 J.W. Murray et al. / Estuarine, Coastal and Shelf Science 58 (2003) 677–697

may provide new analogues. Loch Etive in westernScotland proves to be such an environment.

Loch Etive is a narrow inlet of the sea (fjord)branching off the Firth of Lorn, Scotland (Fig. 1). Onedefinition of a fjord is ‘‘a deep, high-latitude estuarywhich has been. . .excavated or modified by land-basedice’’ (Syvitski et al., 1987). Loch Etive conforms to thisdefinition since it is glacially overdeepened and steep-sided due to erosion during the Quaternary (Howe et al.,2001). The sediment fill is 30–50 m thick (Jones andBlack, 2001; Howe et al., 2002). Because fjords are sitesof active sediment accumulation they provide an ex-cellent opportunity for the study of benthic boundaryprocesses and for providing a high-resolution historicalrecord of environmental change (whether climatic ordue to human impact).

From the point of view of foraminiferal ecology,Loch Etive presents some interesting features. Themaximum water depth (145 m) is comparable to thatof the outer continental shelf yet it occurs in a basinwhich is a mere 2 km wide. Like many other fjords, thedeep water of the Loch is isolated from the open sea bya succession of shallow sills. Major questions to considerare what is the nature of the foraminiferal fauna andhow does it compares with that of the adjacent opencontinental shelf and other silled fjords? Among themacrofauna there are some arctic taxa which reach theirsouthern limit here (see below); does this apply to anyforaminifera? The aim of this preliminary study was toaddress these questions.

2. Area description

Loch Etive has a shallow sill at the entrance (7 m) anda series of five other sills ranging in depth from 4 to 24 m(Edwards and Sharples, 1986). There are two deeperbasins—an outer one (maximum depth �70 m, area

11.35 km2) and an inner one (sill depth 13 m, maximumdepth 145 m, area 16.94 km2) which is the present studyarea. In the deepest area sampled, the surrounding basinslopes are 5–15� (Howe et al., 2001). The mean springtidal range is 3.2 m and surface currents are fast in theentrance to the fjord system (334–399 cm/s) and up to165 cm/s at the narrows between the two main basins(Gage, 1972). There is no information on tidal currentsin deep water but they are likely to be sluggish. Thisfjord has the largest freshwater input of any Scottish sealoch (Hiscock, 1998). The larger part of the catchmentarea drains through the river Awe, which enters the lochat the Bonawe sill. The other part of the catchment area(only slightly smaller) enters the upper basin mainlythrough the river Etive, but also through the Kinglassand other lesser rivers. A hydrographic profile for wintershows inner basin salinities of �22 at the surface (but itcan fall to 1 after rainfall) to 28 at the bottom witha halocline at around 20 m; bottom temperatures rangefrom around 8 to 10 �C throughout the year (Gage,1972). Replenishment of deep waters in the inner basinis episodic (mean 1.3 yr, Edwards and Edelsten, 1977;possibly as infrequent as 2–3 yr, Jones and Black, 2001).Although Gage recorded dissolved oxygen values of40%, depletion becomes greater during prolongedperiods of non-renewal of bottom waters and fell to<20% (<1.5 mg l�1, Jones and Black, 2001) in 2000.The value measured was 0.84 mg l�1 by Winklertitration at the bottom of the Bonawe deep in the April2002, Overnell, pers. comm.), however, anoxia has neverbeen recorded. During periods when the bottom water isstagnant (in the sense of not being replenished) there isreturn of reduced manganese to the sediment. High ratesof total sulphate reduction are associated with high ratesof formation of acid volatile sulphide, which is buried inthe sediments (Overnell et al., 1996; Overnell, 2002). Theorganic content of the sediment (measured as loss onignition) is around 15% but as most of it is resistant

Fig. 1. Map to show the bathymetry of Loch Etive (based on Edwards and Edelsten, 1977; Howe et al., 2001) and sample transect.

679J.W. Murray et al. / Estuarine, Coastal and Shelf Science 58 (2003) 677–697

to decomposition, there is poor correlation betweenthis and oxygen consumption and sulphate reduction(Overnell et al., 1996). From measurements of the ratesof oxygen consumption and inorganic nutrient genera-tion during a stagnant period, Edwards and Grantham(1986) concluded that there was a strong silicate source.

The inner basin is floored with fine grained sediments(63–92% particles finer than 60 lm, Gage, 1972), mostprobably sourced from the River Etive at the head of thefjord, and according to Jones and Black (2001) the rate ofsediment accumulation is 0.7 cm yr�1 (but see thediscussion below). Observations on the sedimentation ofdetritus over a one-year period showed that most of theorganic material is terrestrial in origin and the rest is fromphytoplankton. In deeper waters there was short-termresuspension and redeposition of fine grained material intransit from shallow to deeper levels (Ansell, 1974;Solorzano, 1977). The concentration of bromine in thesurface sediments of the inner basin (400–600 ppm) iscorrelated with the C :N ratio (12–14) and indicates thataround 30–40% of the organic matter is of marine origin(Malcolm and Price, 1984). The production of phyto-plankton varies seasonally with peaks in the spring and itsabundance is controlled not only by the availability ofnutrients but also by hydrographic factors includingfreshwater runoff (Solorzano and Ehrlich, 1977a,b).Primary production was estimated at 70 g C m�2 yr�1 inthat part of the outer basin closest to the sampling areaand light was considered to be the major limiting factor(Wood et al., 1973; there are no data for the inner basin).

The macrofauna in the inner basin is dominated by theophiuroid Amphiura chiajei, five species of polychaetes,four species of bivalves and a penatulid (Gage, 1972),essentially an A. chiajei subcommunity like that of theNorth Sea (Buchanan, 1963) but also similar to the deepmud community of Loch Linnhe and Loch Eil (Pearson,1970). A benthic video survey made during August 1999when the bottom water oxygen concentration was 2.3 mgl�1, showed evidence for a healthy burrowing infauna ofworms and crustaceans (L.Nickell, DunstaffnageMarineLaboratory, pers. comm. cited in Overnell, 2002). Coldwater elements which reach their southern limits in thisarea include a boreo-arctic ascidian (Millar, 1988) andThyasira gouldi (Phillipi), a pan-arctic bivalve (Blacknelland Ansell, 1975). Symbiotic bacteria in the gills of thelatter appear to be sulphur oxidisers (Southward, 1986).

With the exception of a record of presumed liveCrithionina gramen in a macrofaunal study of LochNevis (>1.3 mm, McIntyre, 1961), no studies of livingforaminiferal assemblages from Scottish fjords havepreviously been undertaken and the only previous studyof the dead foraminiferal faunas of Loch Etive wasa brief note in Howe et al. (2002), recording the dom-inance of agglutinated forms in the surface sediment.There have been few studies of the foraminiferal faunasof west Scotland (Williamson, 1858; Heron-Allen and

Earland, 1916; Edwards, 1982; Hannah and Rogerson,1997; Murray, 1985, 2003a,b).

3. Material and methods

Sampling took place at six stations between 26 and138 m water depth (Fig. 1) on 15 and 16 September1998. One short core was collected per station usinga Craib corer, which has a hydraulic damper and a ballclosing system, and takes a core 10–15 cm long and5.7 cm in diameter (Craib, 1965). Additionally, one longcore (EL1, 90 cm) was taken at the deeper station witha controlled-descent gravity corer having a diameter of10 cm, which returned cores with an undisturbedsediment/water interface (Table 1). All cores weresectioned within a short time (hours) of collection andthe samples frozen. With the exception of core E1, thetop centimetres of the Craib cores were sampled in twosections: 0.0–0.5 and 0.5–1.0 cm. For E1, three sectionswere taken: 0.0–0.25, 0.25–0.5, and 0.5–1.0 cm. Belowthis, the cores were sectioned into 1 cm slices down to10 cm. Prior to processing, the samples were thawed in70% ethanol, and then washed on a 63 lm sieve, the>63 lm fraction was stained in rose Bengal (1 g l�1 ofwater) for at least 1 h, washed again on the 63 lm sieveto remove the excess stain, and dried at 50 �C. The longcore samples (2 cm slices throughout) were frozen,weighed wet, freeze dried, weighed dry and five sub-samples for faunal analyses were processed in the sameway but without rose Bengal staining. The justificationfor drying the samples is that although some fragileagglutinated species are destroyed (e.g., Leptohalysis),such forms would not be well preserved in the fossilrecord; the aim of the paper is to help interpret the fossilrecord. The water content was calculated as percent ofwet sample, and the total organic carbon (TOC) contentwas determined by the Leco combustion method (LecoIndustrial Furnace). Salinity of the water overlying thesurface sediments in the short cores was measured withan Atago hand-held refractometer.

Most stained and at least 250 unstained foraminiferawere picked from sub-samples in the top 2 cm. Belowthis, selected samples were analysed for the short coresand five from the long core, EL1. Taxonomy is based on

Table 1

Details of sample locations

Sta. Lat N Long W Depth (m) Equipment

E1 56 28.185 05 09.896 130 Craib corer

EL1 56 28.185 05 09.896 137 Long corer

E2 56 27.859 05 10.898 26 Craib corer

E3 56 27.847 05 10.902 54 Craib corer

E4 56 27.847 05 10.871 72 Craib corer

E5 56 17.846 05 10.790 93 Craib corer

E6 56 17.439 05 10.969 141 Craib corer

E7 56 17.439 05 10.969 138 Craib corer


Murray (2003a). Species diversity was calculated forsamples with >100 individuals using the Fisher alphaindex (Fisher et al., 1943) and the information function,H(S). Cluster analysis, non-metric multidimensionalscaling plots (MDS), and H(S) were determined usingthe program PRIMER v5 (Clarke and Warwick, 1994;Clarke and Gorley, 2001). For the cluster and MDS, thedata were not transformed and similarities were cal-culated using the Bray–Curtis index (Bray and Curtis,1957). All thecamoebians encountered while picking theforaminifera were also picked and determined using thetaxonomy of Scott et al. (2001).

Core E1 was dated via the 137Cs and 210Pb datingmethods (e.g., Cundy and Croudace, 1996). Core sub-samples were counted for 24 h on a Canberra 30%P-typeHPGegamma ray spectrometer to determine the activitiesof 137Cs and other gamma emitters. Limits of detectionwere typically <5 Bq/kg and errors were ca. 20%. 210Pbactivitywas determined by a proxymethod through alphaspectrometric measurement of its grand daughter nuclide210Po. The method employed is based on Flynn (1968),using double acid leaching of the sediment with 209Po asan isotopic tracer andautodeposition of thePo isotopes inthe leachate on to silver discs. Discs were counted ona Canberra 7401 Alpha Spectrometry system for at least150,000 s. Detection limits are 0.1 Bq/kg and errors wereless than 5%. The 210Pbexcess activity was estimated bysubtraction of the value of constant 210Pb activity at depth(0.035 Bq/g), using activity values from a parallel core.Ages were determined using the simple model of 210Pbdating (see Appleby and Oldfield (1992) for review ofdating models).

4. Radiometric dates

137Cs activity shows a slight increase with depth,from surface activities of ca. 100–190 Bq/kg at �10 cm

(Fig. 2). The activities observed are similar to thosefound in surface and near-surface sediments in earlierstudies (Ridgeway and Price, 1987; Williams et al.,1988). 134Cs was not detected in any of the samplesanalysed. Given that the main source of supply of 137Csto Loch Etive is liquid effluent discharge from the BNFLSellafield facility (Williams et al., 1988), the up-coredecline in 137Cs activity observed here most probablyreflects decreasing inputs of 137Cs from Sellafield overtime, due to reductions in 137Cs discharges since themid-1970s. While it is not possible to accurately matchthe observed 137Cs profile to the Sellafield dischargehistory due to the relatively short core length, the down-core increase of 137Cs and the lack of a clear subsurfacemaximum in activity indicates that the sediment samplespost-date the period of maximum discharge fromSellafield in 1975. Assuming that the entire cored depthof sediment is post-1975 gives a minimum sedimentaccumulation rate of 4 mm/yr.

210Pbexcess shows a rapid decline in activity over thecored depth, from 0.33 Bq/g at the sediment surface to0.20 Bq/g at �10 cm depth (Fig. 2). These values of210Pbexcess are similar to those found by Williams et al.(1988) in organic-rich sediments from the inner loch.Applying the simple model of 210Pb dating indicatesa sediment accumulation rate of 0.1 g cm2 yr�1, or 5 mmyr�1 (+/� 1 mm/yr). This is broadly consistent with the137Cs data, and agrees well with sediment accumulationrates of 5–8 mm yr�1 reported in earlier studies (seeWilliams et al., 1988; Howe et al., 2002; Overnell, 2002).It is however, considerably lower than the rates in excessof 1.2 cm yr�1 reported by Ridgeway and Price (1987)for an adjacent inner loch site. This may indicate localvariability in sediment accumulation rate, or more likelyis an artefact of the method used by Ridgeway andPrice, who date the sediments according to the firstappearance of Sellafield-derived 137Cs in the sedi-ment column (corresponding to 1952). This provides

Fig. 2. 137Cs and 210Pbexcess activity vs. depth, Loch Etive core E1. A chronology derived from 210Pb dating is also shown (see text for discussion). All

error bars are 2r (error bars on the 210Pbexcess graph are smaller than the diamond marker symbol used).


a maximum rate of sediment accumulation, as down-ward diffusion of 137Cs may cause pre-1952 sediments tobe labelled with 137Cs, thus giving an erroneously highaccretion rate (e.g., Cundy and Croudace, 1996). The210Pb data presented here indicate that sediment at thebase of the core (i.e. at �10 cm depth) was deposited in1978 (+/� 5 yr, Fig. 2). The consistent decline in 210Pbactivity with depth indicates little sediment homogeni-sation by physical mixing and bioturbation, in agree-ment with earlier studies (see Williams et al., 1988).

5. Sediments

The sediment was very soft and had water content of76.0–86.5%, mean 78.5% in long core EL1, 0–90 cm(n ¼ 31). They are mainly of clay grade and are�packaged� as faecal pellets. TOC from long core EL1(n ¼ 15) yielded values of 5.0% at 0–2 cm, 5.3–5.8%from 2–66 cm, and 6.4–8.0% down to 90 cm. Theparticulate organic matter is primarily terrestrial(spores, wood, cuticle; Ian Harding, pers. comm.).

6. Salinity measurements

At the time of sampling, the salinity of the wateroverlying the core tops ranged from 27 to 30. Thus allstations were brackish and all were from below thehalocline.

7. Foraminiferal results

7.1. Living assemblages

The number of stained individuals in each sampledown to 2 cm was low and below this they were rare ifpresent at all. The standing crop, based on the 0–1 cminterval, ranges from 7 to 35 per 10 cm3 sediment and,on the 0–2 cm interval, 5–36 per 10 cm3 sedimentimplying that the highest abundance consistently wasfound in the surface 0–1 cm (Tables 2 and A1). Asopposed to the other species, Leptohalysis scottii andReophax fusiformis were generally equally abundant atall core depths down to 2 cm and dominated the 1–2 cminterval. The highest values occur at 54 and 130 m andthere is no correlation between standing crop and waterdepth. Species diversity values for the top 0–2 cm arelow: Fisher alpha 1.5–4.7 and H(S) 1.23–1.85. Agglu-tinated foraminifera are dominant (43% at 26 m,otherwise 73–97%). Eggerelloides scaber is commonthroughout (10–46%) but especially at depths >50 mwhere it is the dominant form except at 138 m. Reophaxfusiformis is common (14–53%) except at 26 m anddominant at 138 m. Leptohalysis scottii is fragile and

under-represented in the samples because they weredried during processing. Nevertheless, it reaches abun-dances of 8–27% at depths >50 m. Other agglutinatedtaxa attaining sporadic abundances of 10% includeDeuterammina rotaliformis, Psammosphaera bowmaniand Paratrochammina (Lepidoparatrochammina) spp.The only calcareous forms that are common areElphidium excavatum at 26 and 72 m and Lamarckinahaliotidea at 26 m. On a dendrogram, the samples at 54and 93–138 m form cluster 1 with 60% similarity whilethe two samples with calcareous forms are separateclusters 2 and 3 (Fig. 3). On an MDS plot, cluster 1 isseen to be dominated by E. scaber and R. fusiformis,cluster 2 by E. excavatum and L. haliotidea, whereascluster 3 has no single dominant species.

7.2. Dead assemblages

7.2.1. 0–1 cmThe number of tests per 10 cm3 of sediment shows

a steady decrease with water depth, from 650 at 26 m to67 at 138 m and 53 at 130 m with a major changebetween 54 m (547 tests) and 72 m (189 tests) (Tables 3and A.2). Species diversity is moderately low (Fisheralpha 2.0–3.9, H(S) 0.83–1.73) with the highest values at130 m. The proportion of agglutinated tests is highthroughout (�96% except at 130 m where it is 91–96%).Calcareous tests commonly show evidence of dissolu-tion. Eggerelloides scaber is abundant throughout and isdominant from 26 to 93 m; then Leptohalysis scottii isdominant at 130 m andReophax fusiformis at 130–138 m.Few other species exceed 5% abundance (Ammotiumsp. at 54 m, Deuterammina rotaliformis at 26 m,Miliammina fusca at 26, 93 and 138 m, Trochamminaspp. at 26 m and Elphidium excavatum at 130 m).

Table 2

Summary of live (stained) distributions

Water depth (m) 26 54 72 93 130 138

Core E2 E3 E4 E5 E1 E7

Deuterammina rotaliformis 10 0 0 0 0 1

Eggerelloides scaber 10 46 27 40 34 28

Paratrochammina (L.) spp. 10 1 0 0 0 0

Leptohalysis scottii 0 8 18 17 27 9

Psammosphaera bowmani 5 10 5 0 0 3

Reophax fusiformis 5 14 23 24 26 53

Elphidium excavatum 43 4 23 1 2 5

Lamarckina haliotidea 14 0 0 0 0 0

Fisher alpha 4.7 4.4 2.7 3.3 2.1 1.5

Information function H(S ) 1.75 1.85 1.62 1.65 1.49 1.23

% Agglutinated 43 93 73 89 97 95

0–2 cm, Standing crop

per 10 cm312 36 5 19 X 17

0–1 cm, Standing crop

per 10 cm313 35 7 23 35 21

Species abundances as % for the 0–2 cm interval. X, no sample at

1–2 cm.


Fig. 3. Cluster analysis and MDS plots for live data from 0 to 2 cm. The size of the bubbles on the MDS plots reflects the relative importance of the

named species. The three clusters are labelled 1–3.

Cluster analysis of the 0.5 cm slice data gives twogroups: no. 1 includes the samples from 26 to 93 m andno. 2 those from 130 to 138 m. A similar pattern isshown by the MDS ordination (Fig. 4). The bubbleplots show that several species contribute to cluster 1and that E. scaber is unimportant in cluster 2. SampleE1a is dominated by L. scottii.

7.2.2. Short coresThe dead assemblages from 26 to 93 m are dominated

by Eggerelloides scaber with a range of minor speciesmaking up >5%, the most consistent being Reophaxfusiformis and Elphidium albiumbilicatum (Table 4). At138 m, R. fusiformis is dominant with E. scaber sub-dominant. Agglutinated forms are important through-out. Species diversity is low except at 138 m where it

rises to Fisher alpha 4.8. Although in each core there issome variation in the number of tests per 10 cm3

sediment, the mean values follow the same decrease withwater depth as the surface 0–1 cm samples (Table 2).Small, juvenile, planktonic foraminifera are presentfrom 2 to 5 cm in core E7 at 138 m water depth.

7.2.3. Long core, 0–90 cm, 138 m water depthThe top 10 cm of this core overlap the equivalent

succession of E7. The five samples from 18 to 90 cmhave higher species diversity (Fisher alpha 5.3–9.5, H(S)1.70–2.65) and the proportion of agglutinated formsfalls to 43% at 88–90 cm (Table A2). The number oftests per cm3 sediment is comparable with those at thetop of core E7 but much lower than the number from2 to 10 cm. The dominance of Eggerelloides scaber


Table 3

Summary of dead data for 0–1 cm

Water depth (m)26 54 72 93 130 138

Core E2 E2 E3 E3 E4 E4 E5 E5 E1 E1 E1 E7 E7

Depth (cm) 0.0–0.5 0.5–1.0 0.0–0.5 0.5–1.0 0.0–0.5 0.5–1.0 0.0–0.5 0.5–1.0 0.0–0.25 0.25–0.5 0.5–1.0 0.0–0.5 0.5–1.0

Ammotium sp. 0 0 7 3 0 0 0 0 0 0 0 0 0

Deuterammina rotaliformis 8 6 0 0 0 0 0 0 0 0 0 3 0

Eggerelloides scaber 65 66 57 71 75 82 67 64 26 39 28 35 29

Leptohalysis scottii 1 0 2 0 3 1 3 1 29 6 4 1 2

Miliammina fusca 4 5 4 3 3 0 6 4 3 4 2 1 7

Reophax fusiformis 14 7 8 7 11 8 12 18 26 41 44 47 55

Trochammina spp. 3 5 2 1 1 0 0 1 3 0 0 1 1

Elphidium excavatum 0 0 1 1 2 0 2 1 6 2 6 3 2

Fisher alpha 2.0 2.6 3.9 3.4 3.4 2.8 2.9 3.6 3.9 3.2 3.1 3.2 2.4

Information function H(S) 1.25 1.32 1.65 1.23 1.06 0.83 1.27 1.30 1.73 1.43 1.55 1.39 1.24

% Agglutinated 100 99 99 99 97 96 98 97 91 96 94 97 98

0–1 cm, Tests per 10 cm3 650 547 189 116 53 67

Species abundances in %.

continues down to 32–34 cm and below this Reophaxfusiformis takes over until 64–66 cm. The bottomsample, 88–90 cm, has no strongly dominant species;the most abundant forms in rank order are Stainforthiafusiformis, Elphidium excavatum, Elphidium albiumbili-catum, R. fusiformis and E. scaber. Cluster analysisshows a surface group (cluster A) 0.0–1.0 cm, a broadcluster B of samples down to 48–50 cm but excluding32–34 cm which groups with 64–66 cm in cluster C.Finally, the bottom sample from 88 to 90 cm remainsa separate cluster D. On the MDS plot, the groupingsare the same but it can be seen that 48–50 cm lies in anintermediate position between the main group ofsamples in cluster B and cluster C (Fig. 5). The bubbleplots show that E. scaber and R. fusiformis are im-portant in cluster B and calcareous forms in cluster D.

8. Thecamoebians

Dead individuals were recorded in small numbers inall cores at all depths sampled (Table A2).

9. Discussion

The strong dominance of agglutinated forms with anorganic cement is the outstanding feature of both thelive and dead foraminiferal assemblages of Loch Etive.This resembles some southern Norwegian fjord assemb-lages (Alve and Nagy, 1986; Alve, 1995) but makes themquite unlike any estuarine assemblages previously de-scribed from around the British Isles, which nor-mally are highly calcareous, as discussed below. On aglobal basis, living assemblages dominated by organiccemented agglutinated foraminifera occur to a limitedextent in parts of brackish estuaries, fjords, and shallowbays or lagoons but they are mainly confined to tidal

marshes and to those parts of the ocean deeper than theCCD. It has been argued that some of the fossilagglutinated assemblages must be secondary concen-trations due to (taphonomic) loss of calcareous formsthrough dissolution (Murray and Alve, 1999a,b). Theoccurrence in Loch Etive of primary living agglutinatedassemblages with only a minor calcareous component ismost likely related to the unusual oceanographicconditions in the area. These same conditions causea further postmortem concentration of agglutinatedforms in the dead assemblages through the dissolutionof most of the calcareous tests.

9.1. Influence of unusual oceanographic conditions

The stratification of the water masses with a haloclineat �20 m leads to episodic exchange of bottom water inthe inner basin resulting in the presence of brackishwater of almost constant salinity (28) for periods whichmay exceed one year. In effect, there is no tidal influenceon salinity in the deep waters of this basin. Likewise, theannual variation in bottom temperature is around 2 �C(range 8–10 �C) so the seasonal influence is small too.Thus, the environment is very stable for long periodsalthough there is probably seasonal variation in the fluxof organic material. The episodic variation in dissolvedoxygen does not extend to values low enough to directlyinfluence the benthic foraminifera (from other studies itis apparent that only levels <1 ml l�1 in the bottomwater are critical, e.g., Bernhard et al., 1997). Duringstagnation the dissolved oxygen levels in Loch Etiveare sufficiently low to affect biogeochemical recycling(Edwards and Grantham, 1986; Overnell et al., 1995,1996) and this almost certainly influences the benthos.The most obvious effect may be on the corrosivity of thewater with respect to CaCO3. For instance, althoughophiuroids are common elements of the macrofauna,


Fig. 4. Cluster analysis and MDS plots for dead data from a 0.0–0.5 and b 0.5–1.0 cm samples. For E1, a 0.0–0.25, b 0.25–0.5, c 0.5–1.0 cm samples.

The size of the bubbles on the MDS plots reflects the relative importance of the named species.

their ossicles are not very common in the sediment.Some of the calcareous tests of dead foraminifera showevidence of corrosion and are much reduced in abun-dance in comparison with the living assemblages.However, down-core there are levels with higher abun-dances of calcareous forms (e.g., long core 48–90 cm) sothere has clearly been temporal variation in corrosivityover a longer time period.

In comparison, deeps on the continental shelf west ofScotland also have a similar temperature range andprobably a seasonal flux of organic matter but thesalinity is fully marine throughout the year (34.5–35.2).Oxygen levels are high and there is no evidence ofwidespread carbonate dissolution. Indeed, the sedimentsare rich in carbonate bioclasts and most of the for-aminiferal fauna is hyaline (Murray, 2003b).

Reduction within the sediment of MnO2 derived fromthe water column will be accompanied by an increase inpH, although the magnitude of the increase is difficult toestimate (Overnell, pers. comm.). Oxidation of organiccarbon is bacterially mediated. The role of oxygen islargely to oxidize the reduced intermediates liberated bythe oxidation of organic compounds (nitrite, ammonia,Mn(II), sulphides, low molecular organic acids, etc.). Atthe deep station in the inner basin oxidizing equivalentsfrom the input manganese oxides probably amounted to20–30% of the direct oxygen uptake. At the Airds Baystation oxygen was the dominant electron acceptor(Overnell, 2002). The question arises as to whetherduring these processes the localised release of CO2 inmicroenvironments around the bacterial degradation isresponsible for creating the mildly acidic conditions that


Table 4

Summary of down-core dead assemblage distributions

Core E2 E3 E4 E5 E7

Water depth (m) 26 54 72 93 138

Sediment thickness (cm) 10 10 8 7 10

Eggerelloides scaber Dom Dom Dom Dom Sub-dom

Ammotium salsum M

Leptohalysis scottii M

Miliammina fusca M M

Paratrochammina (L.) spp. M

Reophax fusiformis M M M M Dom

Trochammina spp. M M

Elphidium albiumbilicatum M M M M

Elphidium excavatum M M

Miliolinella subrotunda M M

Fisher alpha 2.0–4.0 2.1–4.0 2.3–3.4 1.9–4.0 2.4–4.8

Information function H(S) 1.40–1.60 1.40–1.60 0.69–2.09 1.23–1.54 1.24–1.92

% Agglutinated 82–100 87–99 87–99 85–90 79–98

Number of tests 10 cm3 sediment 147–1029 421–776 181–632 116–206 67–154

Mean number of tests 10 cm3 sediment 661 500 383 148 115

Dom, dominant, �55%; M, minor, �5%.

cause the carbonate dissolution. Understanding carbon-ate dissolution continues to be a problem. ‘‘A majordifficulty in understanding the dissolution kinetics ofcalcite in seawater is that the saturation state of seawateris generally greater than 0.7 with respect to calcite whichis approximately a pH of only 0.2. Thus, an un-derstanding of the dissolution behavior of calcite inthe ocean, and influences of factors such as inhibitorsand temperature, must be obtained over a pH range ofless than 0.2’’ (Morse and Arvidson, 2002). They alsopoint out that there is a long way to go to bridge the gapbetween carefully controlled laboratory experiments andobservations in complex natural marine environments.

9.2. Features of the living assemblages

Open shelf living assemblages commonly have Fisheralpha species diversity values >5 and informationfunction (H(S)) >0.75 (Murray, 1991). The valuesrecorded for the live assemblages here are lower thanFisher alpha 5 but within the shelf sea range for theinformation function; however, the alpha values arecomparable with those of the southern North Sea(Murray, 1992). The standing crops in Loch Etive arelow (<75 per 10 cm3 at all but one station) compared withcontinental shelves which commonly have >100 individ-uals per 10 cm3 sediment. Stanton Deep on the Scottishshelf has anorganic fluxof 75 gCm�2 yr�1) anda standingcrop (817 per 10 cm3 sediment for 0–1 cm; Murray,2003b). Theorganic flux inEtive ranges from28.4 gCm�2

yr�1 at 26m to 8.7 g Cm�2 yr�1 at 138m (calculated froma primary production of 70 g C m�2 yr�1 using theequation of Berger et al., 1988). Therefore, lower standingcrop values are to be expected.

Both the dominant species, Eggerelloides scaber andReophax fusiformis, are infaunal (Murray, 2003a) butnone of the infaunal species were found to live far belowthe sediment–water interface; virtually all were in thetop 2 cm and 68–84 % in the surface 1 cm. The TROXmodel was put forward by Jorrisen et al. (1995) toexplain depth of life in the sediment and the controlexerted by oxygen and organic flux (given in relativeterms as oligotrophic to eutrophic). Recently, Fontanieret al. (2002) described a transect of stations across theBay of Biscay margin and for the first time provideddata on the values of organic flux in the eutrophic andmesotrophic fields. Their stations D and B are mostsimilar to Etive and both lie in the eutrophic field. D hasan organic flux of 34.1 g C m�2 yr�1 of which the labilefraction is 26.6 g C m�2 yr�1; the oxygen profile reacheszero at �8 mm. Nevertheless, live foraminifera extenddown to 7–8 cm below the sediment surface althoughthey are rare below 4 cm. B has an organic flux of 9.2 gC m�2 yr�1 of which the labile fraction is 6.6 g C m�2

yr�1; the oxygen profile reaches zero at �19 mm.Nevertheless, live foraminifera extend down to 6 cmbelow the sediment surface although they are again rarebelow 4 cm. In Loch Etive, in spite of the high content oforganic matter in the sediment and the relativelylowered oxygen levels in the overlying water, the redoxboundary is a few cm below the sediment surface (basedon geochemical criteria, Overnell et al., 1996) and yet theinfaunal foraminifera are confined to this zone.

Living assemblages dominated by Eggerelloidesscaber are known from Oslofjord and Drammensfjord,Norway (Alve and Nagy, 1986; Alve, 1995), the BalticSea (Lutze et al., 1983) and Arcachon lagoon, France(Le Campion, 1970). On the continental shelf they occur


Fig. 5. Cluster analysis and MDS plots for dead data from core E7 down to 10 cm and long core EL1 from 19 to 90 cm. The sample depths are

expressed by the top number (e.g., 1 ¼ 1:0�2:0 cm). The size of the bubbles on the MDS plots reflects the relative importance of the named species.

in the southern North Sea (Richter, 1967), the CelticSea (Murray, 1979) and the English Channel (Murray,1970). Dead or total assemblages are also knownfrom Gullmar Fjord, Sweden (Qvale et al., 1984) andthe continental shelf off Galicia (Colom, 1984). These as-semblages are associated with salinities >24 for most ofthe year, a wide range of water temperatures (1–20 �C)and substrates of sand or mud (Murray, 1991).

Living assemblages dominated by Reophax fusiformisare known from the Norwegian continental shelf(Mackensen et al., 1985), Celtic Sea (Murray, 1979),

the English Channel (Murray, 1965) and NW Africa(Lutze, 1980). These occur in normal salinities at tem-peratures of �1 to 13 �C and on substrates of sand andmuddy sand and at depths down to >3000 m (Murray,1991). Dead assemblages dominated by this species arerarely recorded perhaps because the tests are relativelyfragile.

Elphidium albiumbilicatum (Fig. 6) has been recordedliving in shallow (e.g., 5–46 m, T 1.6–18.0 �C, salinity 0.1to 31.5, Alve, 1995) and deeper fjordic (e.g., 28–43 m, T3–5 �C, salinity �26, Gustafsson and Nordberg, 1999;


Fig. 6. Elphidium albiumbilicatum (Weiss) SEM pictures. Scale bars 20 lm except for right hand picture, which is 50 lm.

116 m, T 5–8 �C, salinity 34.4–34.8, Gustafsson andNordberg, 2001) as well as in shallow (<6 m) openwaters around the Skagerrak and Kattegat and Alveand Murray (1999) considered that this represented itssouthern limit of distribution. Alve (1995) found it to beone of the most eurythermal and euryhaline species inthe Drammensfjord, Oslofjord. It has not been pre-viously reported from modern sediments in Britishwaters so its occurrence in Loch Etive is of considerableinterest. Apart from temperature control, it was thoughtthat its depth distribution was dependent mainly onsalinity (Alve and Murray, 1999). Brackish salinitiesshowing a narrow range of variation are present in LochEtive throughout the year. This is in marked contrast toBritish tidal estuaries where there is both diurnal andseasonal variation in temperature and salinity.

Leptohalysis scottii seems to be a characteristic fjord-fauna element. In addition to Loch Etive, species of thisvery distinctive, fragile, slender genus are common inScandinavian fjords (e.g., Alve and Nagy, 1986; Alve,2000; Gustafsson and Nordberg, 2000, 2001) as well asin Canadian ones (Blais-Stevens and Patterson, 1998;Patterson et al., 2000). It was particularly abundant(63% of live assemblage and had the third higheststanding crop of all 62 surface samples) at a station inthe inner part of Sandebukta, Oslofjord, which waslocated between the outlet from a paper mill and a river-influenced tidal flat (Alve and Nagy, 1986). Indeed,Blais-Stevens and Patterson (1998, p. 213) speculate that‘‘A high proportion of plant debris, a muddy substrate,and probably lower oxygen levels seem to create idealconditions for Leptohalysis catella’’. It is also knownfrom eastern Canada where it shows a considerableincrease in abundance as a consequence of organicenrichment due to fish farming (Schafer et al., 1995;Scott et al., 2001). Although Ernst et al. (2000), based onexperimental data, regarded L. scottii as being a prom-inent epifaunal/shallow infaunal species intolerant to theenvironmental conditions in deeper sediment layers,Gustafsson and Nordberg (2001) found its maximumabundance sometimes in the 0–1 cm and sometimes inthe 1–2 cm layer and live specimens were present down

to 3 cm (the deepest layer investigated). The trenddescribed by the latter authors is in accordance with ourfindings in Loch Etive.

9.3. Taphonomic impacts

Two taphonomic processes have influenced thecomposition of the dead assemblages: destruction oftests and transport.

The loss of calcareous tests through dissolution isattributed to the unusual oceanographic conditions.Living assemblages which are already rich in aggluti-nated forms give rise to dead assemblages even moredominated by them. Dissolution of calcareous testsclearly takes place soon after death at the surface buttheir greater abundance at depth in some cores indicatesthat this has not always been so intense. It is possible tosimulate natural dissolution of calcareous foraminiferaexperimentally. When normal calcareous-rich originaldead assemblages (ODAs) are treated with dilute acid toremove the calcareous forms the resultant acid-treatedassemblages (ATAs) are composed entirely of organic-cemented agglutinated forms. ATAs from the Celtic Seaand western English Channel have abundant Eggerel-loides scaber at depths of <90 m although the depthrange of the species is down to 150 m (Murray and Alve,2000). Reophax fusiformis is of low abundance in theATAs of the Celtic Sea and western Channel (Murrayand Alve, 2000). Destruction of agglutinated tests mayalso take place. Some taxa are fragile and their tests areprobably under-represented, e.g., Leptohalysis scottiiwhich can be destroyed during sample drying andprocessing. Of the dominant taxa, E. scaber is morerobust than R. fusiformis.

There is evidence of transport of tests both into andwithin Loch Etive. The best indicator of inward trans-port from the open sea is the occurrence of juvenileplanktonic forms in core (E7 2–4 cm). Also, whenbottom water renewal takes place, some live juveniles ofbenthic species may be introduced and some maycolonise the area for short or long periods dependingon the prevailing conditions. The open sea is the


ultimate source of all the benthic foraminiferal colo-nisers (see review by Alve, 1999; Alve and Goldstein,2002). The presence of small numbers of benthic marshtaxa, such as Ammotium sp., Jadammina macrescens andTrochammina inflata, shows that there has been advec-tion of material from the intertidal zone into deeperwaters, even throughout the duration of the long core.This transport is to be expected in such a narrow inletwhere deep water lies close to land. However, becausethe fjord sides are steep, marsh development is verylocalised so there is not a large source area. Whereas inthe Scottish shelf deeps transported foraminiferal testsconstitute a major part of the dead assemblages(�50%), in Loch Etive they are very minor and aremostly from the adjacent marshes.

Thecamoebians are protozoans with unilocular teststhat live in freshwater bogs, lakes and rivers. Someappear to be tolerant of very low salinities (Murray,1967; Scott et al., 2001) but their occurrence in brackishor marine sediments is mainly the result of inwardtransport by rivers. Nevertheless, they can give someindication of the source environment. Centropyxis spp.are tolerant of extreme conditions and occur in coastallakes/ponds affected by salt spray (Scott et al., 2001).Difflugia oblonga lives in gyttja (organic-rich lakesediments composed mainly of plant debris) and isnormally present where the pH is <6.2 (Ellison, 1995).

9.4. Environmental change over the past 200 years

The resolution of down-core faunal changes can beaffected by macrofaunal bioturbation but in Loch Etivethere is no evidence of this on the timescale of the halflife of 210Pb. The down-core foraminiferal recordsprovide information on the sub-recent history of thefjord. The most prominent faunal changes are thereduction (both absolute and relative abundance) inEggerelloides scaber and Reophax fusiformis and theincrease in both Elphidium excavatum and Stainforthiafusiformis towards the base of the long core, EL1(Fig. 7). It can be argued that this is a taphonomicallyinduced pattern. However, if destruction of agglutinatedtests (which no doubt is going on to a certain extent) isthe main reason for the reduction in E. scaber and R.fusiformis, one would expect a much faster reduction inthe latter because it is more fragile. This is not the case.Moreover, if carbonate dissolution is the main reasonwhy S. fusiformis is hardly present at shallower corelevels it should at least be a common component in theliving assemblages but it is not. Unpublished data fromthe Oslofjord, show that in areas where it hardly ispresent in the dead assemblages, it still dominates theliving ones. Consequently, it seems that althoughtaphonomic processes have altered the assemblages,they still contain enough information to suggest that the

environmental conditions in Loch Etive have changedover the time period represented by the core sediments.

The renewal/replenishment of the bottom water ofthe upper basin of Loch Etive is determined largely bythe amount of freshwater on the surface. (After heavyrain the salinity of the surface water can drop to 1.) Atthe sill, the water is partially mixed on the ebb tide, andso during rainy periods the salinity of the incomingwater of the flood tide at the sill is lower than that of thedeep water behind the sill, and tidal exchange affectsonly the surface water. However, this tidal movementdoes very gradually erode the salinity of the deep waterby turbulent diffusion. After a period of prolonged lowrainfall the incoming water is much less brackish andthen water on a flood spring tide is able to displace theold deep water. This renewal of deep water on springtides continues as long as the amount of freshwaterremains low (Overnell, pers. comm.). Thus, higher con-centrations of normal marine foraminifera in the sedi-ment at depth may indicate sustained periods of very lowriver flow during spring tides.

Core EL1 has not been dated, but assuming the samemean accumulation rate as in nearby core E1 (0.5 cm/yr)then it may represent around 180 years of record. At thebase there is a calcareous dominated assemblage witha higher proportion ofmarine species and higher diversity(Fisher alpha >5) as compared to the present-day con-ditions. The upward decrease in Stainforthia fusiformis isparticularly noteworthy. This is an opportunistic speciesflourishing in stressed environments, including oxygendepleted Scandinavian silled fjords (see discussion inAlve, 2003), but it seems to require salinities>28 formostof the year (Gustafsson and Nordberg, 2000). Conse-quently, its increased abundance in the lower parts of thecore suggests somewhat higher salinity than that of thepresent-day. The shallower 48–66 cm core intervalrepresents an intermediate situation with a smallercalcareous component, but still substantially larger thanthe more recent assemblages. This suggests lower cor-rosivity of the bottom/pore water as compared to thepresent-day situation and is probably related to a greatermarine influence. Species diversity is high up to the 18–20cm level which may date from around 1958 and thissuggests that the bottom salinities were closer to that ofnormal seawater during this period than they are now.In core E7 (2–4 cm, representing 1988–1992) another ex-ample of greater penetration of marine water is indicatedby the presence of planktonic tests. Thus the foraminiferalrecord has the potential to provide palaeoceanographicreconstructions in this climatically sensitive fjord.

10. Conclusions

The most outstanding feature of the living benthicforaminiferal assemblages is that they are strongly


Fig. 7. Occurrence of foraminifera in cores E7 and EL1.

dominated by agglutinated foraminifera, a featurenormally associated with deeper water environmentsbelow the CCD or with intertidal marshes. The deadassemblages are further enriched in agglutinated formsindicating that postmortem dissolution of calcareoustests is active. Therefore, this area provides a new ana-logue with which fossil assemblages may be compared.

The cause of these unusual assemblages is attributedto the prevailing oceanographic and sedimentary con-ditions. The isolation of the inner basin of Loch Etive bya series of shallow sills, combined with the high input offreshwater, prevents the continuous exchange of marinebottom water and leads to salinity stratification.Between periods of renewal, the bottom waters becomedepleted in oxygen but they do not become anoxic.Although the sediments are rich in organic matter, mostis terrestrial, and the organic flux is low. Inferredbacterial decay of organic matter, and the geochemicalchanges associated with the oxidation processes, may beresponsible for generating slightly corrosive bottomwaters and causing carbonate dissolution.

The historical record over the past two centuries,interpreted from subsurface foraminiferal assemblages,shows that in the past there was greater renewal of thebottom waters (indicating a lower input of freshwater).Loch Etive has good potential for performing high-resolution palaeoclimatic studies.

Acknowledgements

We thank Julian Overnell, SAMS, and the crew of�Seol Mara� for assistance with sampling. Ian Croudaceis thanked for access to the radiochemistry facilities.Julian Overnell is also thanked for discussion ofaspects of the oceanography and geochemistry andfor very helpful comments on the manuscript. BillAustin (St. Andrews) kindly read the final manuscript.The referees, Dave Scott (Dalhousie, Canada) andJulian Overnell, are thanked for their helpful com-ments.

Appendix Table A

Data on living (st

Water depth (m):138 130

Core E5 E7 E7 E7 E1 E1 E1

Depth (cm) .0 1.0–2.0 0.0–0.5 0.5–1.0 1.0–2.0 0.0–0.25 0.25–0.5 0.5–1.0

Ammotium sp. 0 0 0 0 0 0 0

Cuneata arctica 0 0 0 0 0 0 0

Deuterammina ro 0 1 0 0 0 0 0

Eggerelloides scab 4 15 4 2 10 17 4

Glomospira gordia 0 0 0 0 0 0 0

Paratrochammina 0 0 0 0 0 0 0

Leptohalysis scott 9 1 2 4 2 8 15

Miliammina fusca 0 0 0 0 0 0 0

Psammosphaera b 0 1 1 0 0 0 0

Reophax fusiform 7 14 14 11 11 9 4

Textularia earland 2 0 0 0 1 0 0

Textularia skager 0 0 0 0 0 0 1

Trochammina spp 1 0 0 0 0 0 0

Unidentified aggl 0 0 0 0 0 6 0

Buliminella elegan 0 0 0 0 0 0 1

Elphidium excava 0 2 0 2 0 1 1

Elphidium albium 4 0 0 0 0 0 0

Fissurina margina 0 0 0 0 0 0 0

Lamarckina halio 0 0 0 0 0 0 0

Miliolinella subro 0 0 0 0 0 0 0

Stainforthia fusifo 1 0 0 0 0 0 0

Unidentified calc 0 0 0 0 0 0 0

Number counted 28 34 21 19 24 41 26

Number in sampl 28 34 21 19 24 41 26

Number per 10 c 11 26 16 7 38 64 20

% Agglutinated 82 94 100 89 100 98 92

% Agglut in top 97 99 96

Species abund

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1

ained) foraminifera

26 54 72 93

E2 E2 E2 E3 E3 E3 E4 E4 E4 E5 E5

0.0–0.5 0.5–1.0 1.0–2.0 0.0–0.5 0.5–1.0 1.0–2–0 0.0–0.5 0.5–1.0 1.0–2.0 0.0–0.5 0.5–1

0 0 0 4 0 1 0 0 0 0 0

0 0 0 1 0 0 0 0 0 0 0

taliformis 1 0 1 0 0 0 0 0 0 0 0

er 2 0 0 30 6 1 2 3 1 14 17

lis 1 0 0 2 0 1 0 0 0 0 0

(L.) spp. 1 0 1 1 0 0 0 0 0 0 0

ii 0 0 0 2 1 3 2 1 1 4 2

0 0 0 0 0 0 0 0 0 0 1

owmani 0 1 0 7 1 0 1 0 0 0 0

is 1 0 0 5 2 4 5 0 0 6 8

i 0 0 0 0 0 0 0 0 0 2 0

rakensis 0 0 0 0 0 0 0 0 0 0 0

. 0 0 0 1 0 1 0 0 0 0 0

ut 0 0 0 0 0 0 0 0 0 0 0

tissima 0 0 0 0 0 0 0 0 0 0 0

tum 8 0 1 2 0 1 0 2 3 1 0

bilicatum 0 0 0 2 0 0 0 0 0 1 1

ta 0 0 0 0 0 1 1 0 0 0 0

tidea 2 0 1 0 0 0 0 0 0 0 0

tunda 0 0 0 0 0 0 0 0 0 1 0

rmis 0 0 0 0 0 0 0 0 0 0 0

0 0 0 0 0 0 0 0 0 0 1

16 1 4 57 10 13 11 6 5 29 30

e 32 2 27 57 33 95 11 6 5 29 30

m3 25 2 11 44 26 37 9 5 2 22 23

38 100 50 93 100 85 91 67 40 90 93

1 cm 69 96 79 91

ances as numbers.

Table A2

Data on dead foraminifera

93

E4 E4 E4 E5 E5 E5

0–2.0 3.0–4.0 5.0–6.0 7.0–8.0 0.0–0.5 0.5–1.0 1.0–2.0

0 0 0 0 0 0 0

0 0 0 0 0 0 0

5 3 2 3 3 2 3

0 0 0 0 0 0 0

0 0 0 0 0 0 1

0 1 1 0 1 0 1

1 2 6 0 0 0 0

0 0 0 0 0 0 0

9 178 243 235 86 108 212

0 1 1 0 0 0 0

0 0 0 0 1 0 0

0 0 0 0 0 0 0

0 0 1 1 0 0 0

0 0 0 0 0 0 0

0 0 0 0 0 0 0

2 1 0 0 4 2 5

4 12 10 2 8 6 19

3 10 9 17 0 2 1

0 0 1 0 0 0 0

4 22 26 10 16 30 59

0 0 0 0 0 1 7

0 0 0 0 1 1 0

3 0 1 1 4 3 3

0 0 0 0 0 0 0

0 0 0 0 0 0 0

0 0 2 4 0 1 0

0 0 0 0 3 7 17

0 0 0 0 0 0 0

1 0 0 1 0 0 0

1 1 0 1 0 0 1

0 0 0 0 0 0 0

0 0 0 0 0 0 0

0 0 0 0 0 0 0

0 0 0 0 0 0 0

0 0 0 0 0 0 0

0 0 0 0 0 0 0

0 0 0 0 0 0 0

0 21 24 0 0 0 10

(continued on next page )

691

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Water depth (m) 26 54 72

Core E2 E2 E2 E2 E2 E2 E2 E3 E3 E3 E3 E3 E3 E3 E4 E4 E4

Depth (cm) 0.0–0.5 0.5–1.0 1.0–2.0 3.0–4.0 5.0–6.0 7.0–8.0 9.0–10.0 0.0–0.5 0.5–1.0 1.0–2–0 3.0–4.0 5.0–6.0 7.0–8.0 9.0–10.0 0.0–0.5 0.5–1.0 1.

Ammodiscus

gullmarensis

0 0 0 0 0 0 1 0 0 0 0 2 0 0 0 0

Ammoscalaria runiana 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

Ammoscalaria

pseudospiralis

2 1 0 1 0 1 0 0 0 1 0 0 0 0 1 4

Ammotium sp. 0 0 0 0 0 0 5 16 9 9 2 1 7 1 1 0

Cribrostomoides

jeffreysii

0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

Cuneata arctica 0 0 0 0 0 0 0 1 0 1 1 0 0 0 2 0

Deuterammina

rotaliformis

39 16 21 2 10 17 18 0 0 0 0 0 0 0 0 0

Eggerelloides medius 0 0 0 0 0 0 0 0 2 0 0 0 0 0 0 0

Eggerelloides scaber 319 181 246 160 183 162 157 135 196 195 189 197 244 210 206 173 37

Glomospira gordialis 9 8 4 0 2 4 3 11 8 9 6 1 2 3 2 2

Goesella waddensis 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

Haplphragmoides

wilberti

0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

Jadammina macrescens 0 0 0 0 0 0 0 0 1 0 0 0 0 1 0 0

Lepidotrochammina

ochracea

0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0

Leptohalysis catella 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

Leptohalysis scottii 3 1 3 2 1 1 0 4 1 3 5 2 1 0 7 2

Miliammina fusca 22 15 13 4 13 14 14 10 9 7 6 7 6 6 8 1

Paratrochammina

(L.) spp.

11 13 11 6 43 24 30 5 3 2 10 12 13 14 5 3

Psammosphaera bowmani 0 0 1 0 0 0 0 0 0 0 0 0 1 0 0 1

Reophax fusiformis 67 20 22 48 12 18 15 18 20 11 9 6 6 19 30 16 2

Reophax sp. 0 0 0 0 0 0 0 1 0 0 0 0 0 0 2 0

Spiroplectammina

biformis

2 0 1 0 2 0 0 2 3 2 1 0 0 1 0 0

Textularia earlandi 2 0 0 2 1 0 0 2 1 1 1 1 1 2 0 1

Textularia skagerrakensis 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

Trochammina inflata 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

Trochamminids—

globular

16 14 11 0 2 4 0 5 2 2 8 0 9 6 3 0

Trochammina spp. 0 0 2 0 0 0 0 23 17 25 0 1 0 0 0 0

Webbinella hemisphaerica 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0

Unidentified agglutinated 1 3 2 0 0 1 3 0 0 0 1 2 0 0 1 1

Ammonia beccarii 0 0 1 0 0 2 0 0 0 0 0 0 0 0 1 2

Biloculina inflata 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

Bolivinid 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0

Buccella sp. 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

Bulimina sp. 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

Buliminella elegantissima 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

Cibicides lobatulus 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

Cornuspira involvens 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

Elphidium

albiumbilicatum

0 0 8 23 7 4 4 1 2 2 4 8 4 0 0 6 2

Table A

Water dep 93

Core E4 E4 E4 E4 E5 E5 E5

Depth (cm –1.0 1.0–2.0 3.0–4.0 5.0–6.0 7.0–8.0 0.0–0.5 0.5–1.0 1.0–2.0

Elphidium 0 8 4 3 2 2 1 2

Elphidium 0 0 0 0 0 0 0 1

Elphidium 0 0 0 0 0 0 0 0

Elphidium 0 2 2 0 0 0 0 0

Elphidium 0 0 0 0 0 0 3 19

Epistomin 0 0 0 0 0 0 0 0

Fissurina 0 0 0 0 0 0 0 0

Guttulina 0 0 0 0 0 0 0 0

Haynesina 0 0 0 0 0 0 0 0

Hyalinea 0 0 0 0 0 0 0 0

Lagena su 0 0 0 0 0 0 0 0

Lamarcki 0 0 0 0 0 0 0 2

Miliolinel 0 10 5 0 0 0 0 0

Patellina 0 0 0 0 0 0 0 0

Pyrgo wil 0 0 0 0 0 0 0 0

Quinquelo 0 1 0 0 0 0 0 0

Stainforth 0 0 0 0 0 0 1 2

Triloculin 0 0 0 0 0 0 0 0

Unidentifi 0 2 0 0 0 0 0 0

Number c 2 466 263 330 277 129 168 365

Number i

sample

2 466 1083 1260 1623 129 168 365

Number o

per 10

4 181 421 490 632 100 130.2 142

Number o

% Agglut 6 91 87 92 99 98 97 90

Fisher alp 2.8 3.2 3.2 3.0 2.3 2.9 3.6 4.0

Informati

H(S )

0.83 0.89 1.29 2.09 0.69 1.27 1.30 1.54

Planktoni

Centropyx 1 4 2 1 3

Centropyx 1

Difflugia g 1 1

Difflugia o 5 3 9 4 12 9 6 12

Difflugia p

Difflugia u

Phryganel 6 6 3 5 2 4

Pontigula 1 2

Estimated

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2 (continued )

th (m) 26 54 72

E2 E2 E2 E2 E2 E2 E2 E3 E3 E3 E3 E3 E3 E3 E4 E4

) 0.0–0.5 0.5–1.0 1.0–2.0 3.0–4.0 5.0–6.0 7.0–8.0 9.0–10.0 0.0–0.5 0.5–1.0 1.0–2–0 3.0–4.0 5.0–6.0 7.0–8.0 9.0–10.0 0.0–0.5 0.5

excavatum 0 1 2 1 15 16 2 2 2 1 3 5 1 0 4

magellanicum 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

margaritaceum 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

williamsoni 0 0 0 1 0 1 0 0 0 0 0 0 0 0 0

sp. 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0

ella vitrea 0 0 0 0 0 0 0 0 0 0 0 0 0 0 2

spp. 0 0 1 0 2 0 1 0 0 0 0 1 0 0 0

lactea 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0

germanica 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0

balthica 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

bstriata 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

na haliotidea 0 0 0 0 1 0 3 0 0 0 0 0 0 0 0

la subrotunda 0 0 3 0 34 5 0 0 0 1 3 4 0 0 0

corrugata 0 0 0 0 0 0 0 0 0 0 0 2 0 0 0

liamsoni 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

culina sp. 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0

ia fusiformis 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

a tricarinata 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

ed calcareous 0 0 0 1 0 1 0 0 0 1 0 0 0 0 0

ounted 493 274 353 252 328 276 256 238 276 273 250 250 295 263 275 21

n whole 986 685 2376 378 1968 1840 2645 485 921 1987 1083 1260 1623 1052 275 21

f tests

cm3 sediment

764 531 925 147 766 716 1029 376 714 773 421 490 632 409 213 16

f tests g sediment

inated 100 99 95 89 82 89 96 99 99 98 96 92 98 100 97 9

ha 2.0 2.6 4.0 2.9 3.5 3.7 2.9 3.9 3.4 4.0 3.8 3.8 2.5 2.1 3.4

on function 1.25 1.32 1.31 1.20 1.59 1.60 1.44 1.65 1.23 1.24 1.15 1.04 0.82 0.85 1.06

c

is aculeata 2 1 6 2 2 10 2 3 1 3 4

is constricta

lobulus 1 1 1 1

blonga 8 8 9 6 20 12 15 17 8 11 9 2 12 5

roteiformis

rceolata 3 1

la nidulus 7 2 6 11 7 3 5 6 7 4

sia compressa 3 1 5 4 2 3 1 2 1

year of deposition

Wa

Cor E7 E7 EL1 EL1 EL1 EL1 EL1

Dep –8.0 8.0–9.0 9.0–10.0 18–20 32–34 48–50 64–66 88–90

Am 0 0 0 2 2 0 0

Am 0 1 0 0 0 0 0

Am

p

0 0 1 0 0 0 0

Am 0 0 2 0 0 1 0

Crib 0 0 1 1 0 2 0

Cun 1 0 2 0 2 0 1

Deu

ro

1 2 0 0 0 0 0

Egg 0 0 0 0 0 0 0

Egg 83 78 137 55 34 21 28

Glo 1 2 0 0 0 0 0

Goe 0 0 0 0 0 0 0

Hap 0 0 1 0 0 0 0

Jad 0 0 1 0 2 0 0

Lep

o

0 0 0 0 0 0 0

Lep 0 0 0 1 0 0 0

Lep 16 14 3 5 3 2 3

Mil 13 14 15 9 11 17 15

Par 17 11 2 2 6 2 1

Psa 1 0 0 1 0 0 0

Reo 74 66 104 28 70 24 26

Reo 0 0 0 0 0 0 0

Spir

b

0 0 1 1 0 1 4

Tex 1 2 5 6 4 8 6

Tex 0 0 0 0 0 1 0

Tro 0 0 0 0 0 0 0

Tro

g

0 2 2 4 2 1 4

Tro 0 0 32 19 19 15 26

We 0 0 0 0 0 0 0

Uni 0 0 0 0 0 0 0

Am 0 0 0 0 0 0 3

Bilo 0 0 1 0 0 0 1

Bol 1 1 1 1 0 1 0

Buc 0 0 0 0 2 0 0

Bul 1 0 0 0 0 1 0

Bul 1 0 0 0 0 0 0

Cib 0 0 0 0 0 0 0

Cor 0 0 1 0 2 0 0

Elph 20 32 19 1 19 2 39

Elp 11 11 2 2 30 11 39

(continued on next page )

693

J.W

.Murra

yet

al./Estu

arin

e,Coasta

landShelf

Scien

ce58(2003)677–697

ter depth (m) 93 130 138

e E5 E5 E5 E1 E1 E1 E7 E7 E7 E7 E7 E7 E7 E7 E7

th (cm) 2–3 4–5 6–7 0.0–0.25 0.25–0.5 0.5–1.0 0.0–0.5 0.5–1.0 1.0–2.0 2.0–3.0 3.0–4.0 4.0–5.0 5.0–6.0 6.0–7.0 7.0

modiscus gullmarensis 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

moscalaria runiana 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

moscalaria

seudospiralis

1 1 0 0 0 0 0 0 0 0 0 0 0 0 0

motium sp. 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

rostomoides jeffreysii 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

eata arctica 0 1 0 0 0 1 0 0 0 0 1 0 0 0 0

terammina

taliformis

0 0 1 0 0 0 2 0 9 7 2 1 0 4 1

erelloides medius 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

erelloides scaber 172 150 199 9 19 15 25 29 54 96 61 76 88 79 86

mospira gordialis 2 0 0 0 0 0 0 0 0 2 1 3 2 0 1

sella waddensis 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

lphragmoides wilberti 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

ammina macrescens 0 0 0 0 0 0 1 0 0 0 0 0 1 1 0

idotrochammina

chracea

0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

tohalysis catella 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

tohalysis scottii 2 9 0 10 3 2 1 2 1 4 2 1 4 5 9

iammina fusca 5 7 5 1 2 1 1 7 10 10 12 9 16 12 7

atrochammina (L.) spp. 8 13 6 0 1 2 3 1 0 20 6 9 18 10 18

mmosphaera bowmani 0 0 0 0 0 0 0 0 0 0 3 3 1 1 0

phax fusiformis 34 38 62 9 20 24 34 56 86 112 97 127 109 101 95

phax sp. 0 0 0 0 1 0 0 0 0 1 0 0 0 0 0

oplectammina

iformis

0 0 0 0 0 0 0 0 0 2 0 0 0 0 0

tularia earlandi 0 4 2 1 0 2 2 2 2 1 3 2 1 2 0

tularia skagerrakensis 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

chammina inflata 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0

chamminids—

lobular

21 8 7 1 0 0 1 1 6 2 3 7 4 5 1

chammina spp. 0 1 0 0 1 4 0 0 0 0 0 0 0 0 0

bbinella hemisphaerica 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

dentified agglutinated 0 0 0 1 0 0 0 0 0 3 0 0 1 0 0

monia beccarii 0 0 0 0 0 0 0 0 0 3 1 0 0 0 0

culina inflata 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

ivinid 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0

cella sp. 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

imina sp. 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0

iminella elegantissima 0 0 0 0 0 0 0 0 0 3 0 0 1 0 1

icides lobatulus 0 0 0 0 0 0 0 0 0 0 1 0 1 2 0

nuspira involvens 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

idium albiumbilicatum 19 19 44 0 1 0 0 0 2 8 7 11 11 17 17

hidium excavatum 0 0 3 2 1 3 2 2 15 22 16 17 8 13 7

Table A2 (co

Water depth

Core E7 E7 E7 EL1 EL1 EL1 EL1 EL1

Depth (cm) 0 7.0–8.0 8.0–9.0 9.0–10.0 18–20 32–34 48–50 64–66 88–90

Elphidium m 0 0 0 0 0 0 0 0

Elphidium m 0 0 0 0 2 0 0 2

Elphidium w 0 0 2 2 0 0 2 2

Elphidium sp 0 0 0 0 2 1 1 0

Epistominella 0 0 0 0 1 0 0 0

Fissurina spp 1 0 0 0 1 1 3 2

Guttulina lac 0 0 0 1 2 0 0 0

Haynesina g 0 0 0 0 0 0 0 0

Hyalinea bal 0 0 0 0 0 0 1 0

Lagena subs 0 0 0 0 0 1 0 0

Lamarckina 0 0 0 0 1 0 0 2

Miliolinella s 18 2 2 0 3 8 3 9

Patellina cor 0 1 0 0 0 0 0 0

Pyrgo william 0 0 0 0 1 0 2 1

Quinquelocul 1 1 0 0 1 0 8 1

Stainforthia 0 0 2 0 2 3 5 45

Triloculina t 0 0 0 0 0 0 1 1

Unidentified 0 2 0 0 0 0 1 2

Number cou 263 248 242 336 154 222 137 263

Number in w 395 248 242 336 156 222 137 263

Number of t

10 cm3 sed

116 298 242 74 28 40 30 62

Number of t 76 38 47 34 62

% Agglutina 83 84 79 92 86 69 69 43

Fisher alpha 3.2 4.8 3.8 5.3 9 5.3 9.5 6.4

Information 1.71 1.88 1.92 1.7 2.27 2.23 2.65 2.47

Planktonic

Centropyxis 3 2 13 8 8 11 10 10

Centropyxis

Difflugia glo 3

Difflugia obl 19 10 16 37 28 27 19 66

Difflugia pro 1

Difflugia urc

Phryganella 2 10 14 26 28 17 17

Pontigulasia 5 3 4 1 1 2 1

Estimated ye

deposition

1978 1958 1930 1898 1866 1818

Species

694

J.W

.Murra

yet

al./Estu

arin

e,Coasta

landShelf

Scien

ce58(2003)677–697

ntinued )

(m) 93 130 138

E5 E5 E5 E1 E1 E1 E7 E7 E7 E7 E7 E7 E7 E7

2–3 4–5 6–7 0.0–0.25 0.25–0.5 0.5–1.0 0.0–0.5 0.5–1.0 1.0–2.0 2.0–3.0 3.0–4.0 4.0–5.0 5.0–6.0 6.0–7.

agellanicum 0 0 0 0 0 0 0 0 0 0 0 0 0 0

argaritaceum 0 0 0 0 0 0 0 0 0 0 0 1 0 0

illiamsoni 1 0 0 0 0 0 0 0 0 0 0 1 1 0

. 0 0 0 0 0 0 0 0 0 0 0 0 0 0

vitrea 0 0 0 0 0 0 0 0 0 0 0 0 0 0

. 0 0 0 0 0 0 0 0 0 0 0 0 0 0

tea 0 0 0 0 0 0 0 0 0 0 0 0 0 0

ermanica 0 0 0 0 0 0 0 0 0 0 0 0 0 0

thica 0 0 0 0 0 0 0 0 0 0 0 0 0 0

triata 0 0 0 0 0 0 0 0 0 0 0 2 0 0

haliotidea 0 0 0 0 0 0 0 0 0 0 0 0 0 0

ubrotunda 0 2 2 1 0 0 0 0 2 9 4 5 8 9

rugata 0 0 0 0 0 0 0 0 0 1 0 0 0 0

soni 0 0 0 0 0 0 0 0 0 0 0 0 0 0

ina sp. 0 0 0 0 0 0 0 0 0 0 0 1 0 0

fusiformis 0 0 0 0 0 0 0 0 1 0 0 0 0 0

ricarinata 0 0 0 0 0 0 0 0 0 0 0 0 0 0

calcareous 0 0 0 0 0 0 0 0 1 1 0 1 0 0

nted 265 253 331 35 49 54 72 101 189 308 220 277 275 262

hole sample 530 380 331 35 48 54 72 101 189 308 330 300 367 349

ests per

iment

206 148 129 55 75 42 56 78 74 128 119 143 136 154

ests g sediment

ted 92 92 85 91 96 94 97 98 89 84 87 86 89 84

2.1 2.3 1.9 3.9 3.2 3.1 3.2 2.4 2.9 4.8 4 4.3 4 3.5

functionH(S ) 1.23 1.42 1.23 1.73 1.43 1.55 1.39 1.24 1.56 1.88 1.71 1.69 1.71 1.79

17 12 1

aculeata 1 1 1 1 2 3 8 1 1 3

constricta 1 3

bulus 1 2 2 1

onga 13 8 13 2 1 3 13 32 14 20 14 20

teiformis 1 1

eolata

nidulus 2 3 7 2 1 1 1 2 3 10 12 3 3 10

compressa 2 1 2 4 3 6 1 1 5 2

ar of 1998 1992 1988

abundances as numbers.


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