Polar Biol (2000) 23: 373 382

Springer-Verlag 2000

ORIGINAL PAPER

M. J. Whitehouse J. Priddle M. A. Brandon

Chlorophyll/nutrient characteristics in the water masses to the north of South Georgia, Southern Ocean

Accepted: 27 November 1999

Abstract Chlorophyll a and nutrient concentrations along with temperature and salinity values were measured at 22 CTD stations along a 735-km transect running to the northwest of the island of South Georgia, Southern Ocean. Measurements were repeated during ve summer surveys (January and February 1994, January 1996, December 1996, January 1998) and one spring survey (October 1997). The transect sampled SubAntarctic Zone water in the north, Polar Frontal Zone water and Antarctic Zone water in the south. Chlorophyll a concentrations were lowest to the north of the transect and frequently high (up to 17 mg m)3) in the deep open ocean of the Antarctic Zone. Sub-surface peaks were measured in all zones and chlorophyll a was detectable to a depth of 150 m. There was a clear latitudinal temperature gradient in the near-surface waters (050 m), the warmest water occurring in the north (~12 C), and the coolest in the Antarctic Zone (~2 C). There was also a well-dened latitudinal gradient in summer near-surface silicate concentrations (~2, 4, and 10 mmol m)3 in the Sub-Antarctic Zone, the Polar Frontal Zone and the Antarctic Zone, respectively), increasing to >20 mmol m)3 near South Georgia. Distinct dierences in silicate concentrations were also evident in all three zones to a depth of 500 m. Nearsurface nitrate and phosphate concentrations were relatively low to the north of the transect (~14 and 1 mmol m)3, respectively) and higher in the Polar Frontal Zone and Antarctic Zone (~18 and 1.4 mmol m)3, respectively). Ammonium and nitrite were restricted to the upper 200 m of the water column, and exhibited sub-surface concentration peaks, the lowest being in the Sub-Antarctic Zone (0.68 and 0.25 mmol m)3, respectively) and the highest in the Antarctic Zone (1.72 and 0.29 mmol m)3, respectively). Surface (~6 m) spring nutrient measurements provided
M. J. Whitehouse (&) J. Priddle M. A. Brandon British Antarctic Survey, High Cross, Madingley Road, Cambridge CB3 0ET, UK e-mail: m.whitehouse@bas.ac.uk

an indication of pre-bloom conditions; ammonium and nitrite concentrations were low (~0.27 and 0.28 mmol m)3, respectively), while silicate, nitrate and phosphate concentrations were high and similar to previously measured winter values (e.g. ~26, 23, 2 mmol m)3, respectively in the Antarctic Zone). Although the values measured were very variable, and there was some evidence of a seasonal growth progression, the chlorophyll a and nutrient distribution patterns were dominated by intercruise (interannual) factors. Approximate nutrient depletions (spring minus summer) appeared similar in the Polar Frontal Zone and Antarctic Zone for nitrate and phosphate, while silicate showed a marked latitudinal increase from north to south throughout the transect. Highest chlorophyll a concentrations coincided with the highest apparent silicate depletions over the deep ocean of the Antarctic Zone. In this area, relatively warm, easterly owing Antarctic Circumpolar Current water meets cooler, westerly owing water that is inuenced by the WeddellScotia Conuence and is rich in nutrients, especially silicate.

Introduction
The hydrography associated with the Antarctic Circumpolar Current (ACC) in the vicinity of South Georgia is complex. To the west of the island, the Scotia Ridge diverts its main current north before it resumes an easterly course. The current ow is further complicated by the Maurice Ewing Bank to the northwest of South Georgia, while to the north and east of the island the ACC is met by water inuenced by the Weddell Gyre. The eastward-owing ACC to the north of South Georgia comprises three zones separated by frontal systems (Gordon et al. 1977; Sievers and Nowlin 1988; Orsi et al. 1995; Trathan et al. 1997; Moore et al. 1999). Furthest north is the Sub-Antarctic Zone (SAZ), the

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southern limit of which is dened by the Sub-Antarctic Front (SAF). Southwards from the SAF is the Polar Frontal Zone (PFZ), the Polar Front (PF), and then furthest south, the Antarctic Zone (AAZ). Another front, the Southern ACC Front, is generally located near the poleward extreme of the ACC, close to its boundary with continental waters. At South Georgia the Southern ACC Front, along with Weddell-Scotia Conuence waters, are inected north then west around the eastern and northern anks of the island (Whitworth et al. 1994; Orsi et al. 1995; Trathan et al. 1997; Brandon et al. 1999). The positions of these frontal systems are subject to both large-scale oceanographic and mesoscale variability, with meanders and core-ring shedding recorded for the SAF and the PF (Gordon et al. 1977; Joyce and Patterson 1977; Legeckis 1977; Lutjeharms and Baker 1980; Bryden 1983; Klinck 1985; Nowlin and Klinck 1986). Previous research cruises around South Georgia during the austral summer have recorded great variability in near-surface temperatures, phytoplankton production and nutrient depletion (Hart 1934, 1942; Hardy and Gunther 1935; Clowes 1938; Deacon 1977; Fryxell et al. 1979; El-Sayed and Weber 1982; Ronner et al. 1983; Priddle et al. 1986, 1995; Whitehouse et al. 1993, 1996a, b, 1999). Hart (1934, 1942) described variability in the timing of phytoplankton growth in the Southern Ocean and suggested a latitudinal succession in growth initiation and peak. Latitudinal temperature gradients were identied by Deacon (1977). Hardy and Gunther (1935) and Clowes (1938) documented seasonal phytoplankton utilisation of phosphate and silicate, and also suggested that low silicate concentrations may limit Southern Ocean phytoplankton growth. Whitehouse et al. (1996a) documented chlorophyll a and nutrient concentrations measured around the island that were frequently atypical of the Scotia Sea. For instance, chlorophyll a concentrations >25 mg m)3, and silicate depleted to <1 mmol m)3 have been measured to the northwest of the island. During the Islas Orcadas cruise 17 in 1978, El-Sayed and Weber (1982) recorded chlorophyll a concentrations between 11 and 13 mg m)3 during late winter/early spring (5 October 1978) at their deep-ocean station 34, ~290 km to the north of South Georgia. During the same cruise, Fryxell et al. (1979) speculated that the phytoplankton between stations 34 and 33 (110 km further south) may have been seeded from southeast of South Georgia and carried in a large semi-circular path or possibly entrapped in some sort of gyre. Recent British Antarctic Survey cruises have measured particularly high levels of chlorophyll a in the deep ocean to the north of South Georgia between the island and the PF (Whitehouse et al. 1996b). In this paper we present chlorophyll a, nutrient, temperature and salinity data collected during ve cruises to the north of South Georgia between January 1994 and January 1998. We describe and quantify the extent of variability (e.g. interzonal and intercruise), consider the range of apparent

summer nutrient depletions, and highlight some of the factors that are potentially important for high phytoplankton productivity.

Materials and methods

A standard transect comprising 22 CTD casts sited ~35 km apart was run from 48.0S, 43.8W to 53.9S, 38.7W (Figs. 1, 2). The transect, referred to as the Maurice Ewing Bank transect for the major topographical feature over which it passes, crossed from the SAZ, through the PFZ, and into the AAZ to the northwest of South Georgia. Oceanographic measurements were made from the RRS James Clark Ross between 25 January and 24 February 1994 (cruise JR06 transects T02 and T74), 57 January 1996 (cruise JR11), 1621 December 1996 (cruise JR17), 2325 October 1997 (cruise JR25) and 1721 January 1998 (cruise JR28). During cruise JR06 stations 821 were occupied, and the cruise JR25 transect comprised continuous surface measurements of a non-toxic seawater supply while the ship was underway. All stations were occupied during the other cruises. Water samples were collected at all stations using a General Oceans 12 bottle rosette system along with a Mk IIIB Neil Brown CTD. Standard sample depths were ~6 m (from the ship's nontoxic seawater supply), and 20, 40, 60, 80, 100, 125, 150 and 200 m, and a further four depths sampled between 200 m and the bottom of the water bottle cast. Sub-samples from the water bottles and the ship's non-toxic supply were ltered through a mixed ester membrane (Whatman WME, pore size 0.45 lm), and the ltrate was analysed colorimetrically for dissolved nitrate+nitrite (NO3+NO2-N), nitrite (NO2-N), ammonium (NH4-N), silicate [Si(OH)4-Si] and phosphate (PO4-P) using a Technicon segmented ow analyser (Whitehouse and Woodley 1987; Woodward 1992; Whitehouse 1997;

Fig. 1 Location of the transect stations in relation to the Maurice Ewing Bank, South Georgia, the 500-m and 2000-m isobaths and the area potentially inuenced by Weddell-Scotia Conuence water

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Fig. 2 Transect bathymetry and sampling locations, along with summer chlorophyll a, physical oceanography and nutrient measurements made in the near-surface waters (050 m). The approximate extent of the zones is identied in the silicate panel. The cruise identiers are: n JR06 T02 (25 January 1994), , JR06 T74 (24 February 1994), h JR11 (57 January 1996), s JR17 (1621 December 1996), e JR28 (1721 January 1998) and a median line is tted to all parameters

376 Whitehouse and Preston 1997). Nitrite concentrations, which were low throughout this study, have not been deducted from the combined nitrate+nitrite measurements, which are referred to as nitrate throughout this paper. Phosphate was not analysed during cruise JR06 transect T02, and ammonium was not analysed during cruise JR28. A further water bottle sub-sample was ltered through a glass bre lter (Whatman GF/F, nominal pore size 0.7 lm), and the retained particulate material was analysed for chlorophyll a and phaeopigments (Parsons et al. 1984). Filters were extracted in 90% acetone and the uorescence of the extract measured, before and after acidication, using a Sequoia Turner model 112 uorometer, or a custom-built unit (with emission and excitation lters comparable to those on the Turner instrument). Both instruments were calibrated against commercially prepared chlorophyll a standards (Sigma, St. Louis, Mo.). The software package Minitab 12.1 (Minitab, State College, Pa.) was used to describe and quantify the variability of values measured along the transect. A General Linear Model of nested factors was used to obtain variance component estimates from adjusted sums of squares for cruise, zone and station. The remaining error was assumed to be due to sampling depth.

and PFZ water (Gordon et al. 1977; Sievers and Nowlin 1988), we considered stations 612 to comprise predominantly PFZ water. However, we have labelled stations 15 as the northern zone (NZ) because, although SAZ water was the most prevalent there, there were also obvious incursions of cooler, fresher, higher-nutrient PFZ water. To summarise, we have separated our transect into the NZ (stations 15), the PFZ (stations 612) and the AAZ (stations 1322) (Fig. 1). Near-surface (050 m) summer hydrographic observations The distance between CTD casts (~35 km) did not allow accurate characterisation of front-specic variability; however, there were clear dierences between the three zones (Fig. 2). Average chlorophyll a concentrations were low in the NZ and generally highest in AAZ water (Table 1). Northern PFZ water chlorophyll a values were also low, possibly reecting incursions of SAZ water, whereas the highest PFZ concentrations were found between stations 8 and 12 (median 1.4 mg m)3, maximum 6.2 mg m)3). The peak at station 13 (median 3 mg m)3) may have been related to the PF. Within the AAZ, high chlorophyll a levels were measured at all sites between the PF and South Georgia, with the highest value at station 15 (17 mg m)3). Chlorophyll a levels appeared particularly high over deep water between stations 16 and 19 (median 2.8 mg m)3, maximum 14.3 mg m)3), and again above the South Georgia shelf at stations 21 and 22 (median 4.5 mg m)3, maximum 10.2 mg m)3). Phaeopigment concentrations followed a similar pattern with lowest values found in the NZ and PFZ, and highest values in the AAZ. Concentrations were particularly high in the PFZ at station 8 (median 0.36 mg m)3), and in the AAZ at station 16 (median 0.62 mg m)3), and at stations 21 and 22 over the South Georgia shelf (median 0.67 mg m)3). Temperature and salinity dierences between the three zones were particularly marked, with warmer, more saline water apparent in the NZ (Fig. 2, Table 1). The most northerly three stations were distinctly warmer (median 9.7 C), while cooler water was

Results
Water masses We used standard potential temperature and salinity envelopes previously described for the Scotia Sea to identify SAZ, PFZ and AAZ water, and to locate the position of the SAF and PF on our transect (Gordon et al. 1977). During the six transects the PF was repeatedly found between stations 12 and 13, and was always on the southern ank of the Maurice Ewing Bank. Although this is in contrast to the climatological position for the PF given by Orsi et al. (1995), it is in agreement with the ndings of Gordon et al. (1977), Trathan et al. (1997) and Moore et al. (1999). Therefore, the southern stations 1322 predominantly comprised AAZ water. Within this zone there was evidence of water from the Scotia Sea to the southeast of South Georgia (Trathan et al. 1997), and a distinct water type on the South Georgia shelf at station 22 (Brandon et al., in press). The location of the SAF was less clearly dened in our data set. Using the temperature, salinity and nutrient characteristics previously described for SAZ

Table 1 Average summer (December to February) near-surface (050 m) values Northern zone n Chlorophyll a (mg m)3 ) Phaeopigment (mg m)3) Temperature (C) Salinity Silicate (mmol m)3) Phosphate (mmol m)3) Nitrate (mmol m)3) Ammonium (mmol m)3) Nitrite (mmol m)3) 35 27 29 31 45 45 43 35 45 Median (range) 0.73 0.21 9.28 34.11 2.17 1.07 14.84 0.36 0.19 (0.241.52) (0.010.57) (4.6312.03) (33.8634.68) (0.488.22) (0.431.84) (2.1620.11) (0.081.35) (0.100.30) Polar frontal zone n 76 58 72 72 90 73 88 73 89 Median (range) 1.18 0.21 5.36 33.92 4.29 1.41 18.23 0.39 0.21 (0.116.18) (0.001.04) (2.747.57) (33.7934.06) (1.0119.77) (1.011.83) (10.2423.44) (0.121.56) (0.110.28) Antarctic zone n 120 97 119 121 129 99 125 104 129 Median (range) 2.88 0.38 2.94 33.85 10.10 1.34 17.44 0.59 0.30 (0.3716.98) (0.002.33) (1.014.49) (33.6833.96) (1.0830.41) (0.752.02) (10.0424.20) (0.141.93) (0.220.35)

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evident in the PFZ, with a temperature gradient southwards to the PF. AAZ near-surface temperatures were less variable than those in the other two zones, and the southernmost station on the transect over the South Georgia shelf was sited in the coolest water (median 2.2 C). Although highly variable, the most saline near-surface waters were found in the NZ (generally >34), where particularly high values (median 34.6) were measured during cruise JR17. Near-surface salinity concentrations in the PFZ and the AAZ were similar and fresher than those to the north, and again, the southernmost station 22 over the South Georgia shelf diered from the other AAZ stations (median salinity 33.79). There was a clear latitudinal gradient in silicate concentrations, which were low in the NZ and highest to the south near South Georgia (Fig. 2, Table 1 ). A peak in values at station 6 (median 4.8 mmol m)3) may have been associated with upwelling at the SAF. Although the median silicate concentration in the AAZ was 10.1 mmol m)3, values were very variable, and there was an obvious gradient evident towards the south of the transect. From a peak of 10.6 mmol m)3 at station 14, median concentrations dropped to 7.3 mmol m)3 between stations 15 and 18, before increasing sharply towards South Georgia (median >20 at stations 21 and 22, but note variability). Average near-surface phosphate concentrations were distinctly lower in the NZ compared with the PFZ and AAZ, while variability was lowest in the PFZ. At the mid-AAZ stations 1519, there was a dip in concentrations (median 1.21 mmol m)3), apparently coincident with reduced silicate levels. A similar pattern was evident for nitrate, with near-surface concentrations in the NZ very variable but low compared with the PFZ and the AAZ. Lowest concentrations in the AAZ did not appear to be coincident with reduced silicate or phosphate, but were located further south between stations 18 and 21 (median 15.8 mmol m)3). Near-surface ammonium concentrations were very variable throughout the transect, with the lowest values in the NZ and PFZ. There were high concentrations at stations 14 during cruise JR11 (median 0.74 mmol m)3 at station 2), which appeared to be coincident with cooler, fresher water that contained elevated concentrations of phosphate and nitrate. Levels were also generally high at station 7 in the PFZ (median 0.65 mmol m)3). Overall, concentrations were highest in the AAZ, and appeared elevated at the mid-zone stations 1618 (median 0.84 mmol m)3, maximum 1.93 mmol m)3), possibly coincident with silicate and phosphate depletion and higher concentrations of chlorophyll a. Near-surface nitrite concentrations were low throughout. The lowest values were measured in the NZ and, although PFZ levels were similar, there was a distinct gradient up to higher concentrations at the PF. Concentrations in the AAZ were relatively higher, although lower values were evident near South Georgia (median 0.27 mmol m)3 at station 22).

Mid-water (to ~500 m) summer hydrographic observations The majority of chlorophyll a was found in the upper 50 m of the water column, where there was an indication of a sub-surface maximum in all zones (e.g. median 3.2 mg m)3 at 40 m in the AAZ) (Fig. 3). However, lesser values were evident between 50 and 100 m (at ~60 m 0.54, 1.01, 1.75 mg m)3 in the NZ, PFZ and AAZ, respectively), and measurable traces were found down to 155 m (~0.06 mg m)3 in all zones). The majority of phaeopigment was measured in the upper 100 m of the water column, with an apparent peak at ~5060 m in the NZ and PFZ. Trace phaeopigment concentrations (0.060.14 mg m)3) were evident in all zones to a depth of 200 m presumably as a result of sinking dead cells. The dierent zonal temperature and salinity properties were clearly evident in the vertical proles (Fig. 3). AAZ water was distinctly cooler and showed a characteristic temperature minimum value (median 0.3 C) between the depths of 100 and 150 m. PFZ waters were warmer and those in the NZ warmer still, and they remained so down the water column to below 500 m. The salinity pattern was not so distinctive. NZ surface water was clearly more saline than PFZ and AAZ water to a depth of 150 m, which is about as deep as SAZ surface water reaches (Sievers and Nowlin 1988). Between 150 and 500 m NZ and PFZ waters appeared more alike and were presumably indicative of Antarctic Intermediate Water, as opposed to the Circumpolar Deep Water, which underlies AAZ surface water (Peterson and Whitworth 1989). Silicate concentrations in the three zones were distinct to a depth of at least 500 m (Fig. 3). The AAZ surface water and its underlying Circumpolar Deep Water held by far the highest concentrations with a median of 55.0 mmol m)3 at ~200 m compared with 22.5 and 15.1 mmol m)3 for the corresponding depth in the PFZ and NZ respectively. Interzonal dierences were not so clear for phosphate and nitrate. The surface waters of the AAZ and the PFZ were similar in concentration to a depth of 100 m, and there was some evidence of maximum depletion occurring below the surface layer (deeper than 6 m). There would appear to be higher nutrient levels below the AAZ surface waters compared with those of the PFZ (e.g. at 200 m, 2.21 and 1.99 mmol m)3 phosphate, 31.0 and 25.2 mmol m)3 nitrate respectively). Phosphate and nitrate concentrations were generally lower in the NZ surface waters and their underlying waters (at 200 m, 1.58 and 24.5 mmol m)3 phosphate and nitrate, respectively). However, there would appear to be some agreement between phosphate concentrations in all three zones below a depth of ~400500 m, with a similar parity evident for nitrate. Ammonium and nitrite concentrations, other than trace values, were limited to the upper 200 m of the water column (Fig. 3). There were clearly higher values of ammonium in the AAZ, and all three

378 Fig. 3 Median summer chlorophyll a, physical oceanography and nutrient values for the surface to mid-waters (0500 m) in the three zones

zones, but particularly the AAZ and the PFZ, exhibited sub-surface concentration peaks towards the bottom of the upper mixed layer (UML) AAZ 1.72 mmol m)3 at ~90 m, PFZ 0.99 mmol m)3 at ~60 m, NZ 0.68 mmol m)3 between 65 and 110 m. Nitrite concentrations were generally low but the highest concentrations were clearly in the AAZ where median values of 0.29 mmol m)3 extended from the surface to below 100 m. Minor sub-surface concentration peaks were evident in the PFZ (0.26 mmol m)3 at 90105 m) and the NZ (0.25 mmol m)3 at ~65 m). Surface (~6 m) spring nutrient observations Surface measurements made along the Maurice Ewing Bank transect during the austral spring (2325 October 1997) provided an indication of pre-growing season nutrient conditions, as only a limited amount of phytoplankton growth had occurred at this time. Although median surface chlorophyll a concentrations were ~0.4 and 1.8 mg m)3 in the PFZ and AAZ respectively, phaeopigments were low or absent (median 0.02 and 0 in the PFZ and AAZ), suggesting only recent phytoplankton growth. Furthermore, ammonium and nitrite were only present at low levels (0.27 and 0.28 mmol m)3, respectively), while silicate, nitrate and phosphate concentrations were high, ~26, 23 and 2 mmol m)3, respectively in the AAZ, similar to

Fig. 4 Spring (JR25, 2325 October 1997, pre-bloom) silicate, nitrate and phosphate concentrations measured in the surface waters (~6 m)

previously measured winter values (Whitehouse et al. 1996a). Silicate concentrations showed a clear latitudinal gradient with values <4 mmol m)3 in the NZ rising to ~10 mmol m)3 at the SAF (Fig. 4). In the PFZ, concentrations rose from ~12 mmol m)3 between stations 6 and 10 to ~16 mmol m)3 at station 12, prior to a steep gradient into AAZ waters 25 mmol m)3 in the north of the zone and 33 mmol m)3 near South Georgia. Nitrate and phosphate concentrations were far more uniform. Both were relatively low in the NZ (13 and 1.0 mmol m)3, respectively), but PFZ and AAZ concentrations were comparable. Nitrate concentrations were ~25 mmol m)3 throughout the PFZ, 2324 mmol m)3 in the northern section of the AAZ and ~27 mmol m)3 near to South Georgia. Similarly, phosphate concentrations of ~1.9 mmol m)3 throughout the

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PFZ dipped to ~1.8 mmol m between stations 15 and 18 prior to rising to >2 mmol m)3 near the island.

)3

Discussion
The cause and magnitude of variability along the transect The values measured along the Maurice Ewing Bank transect obviously varied considerably in both time and space. Although all of the CTD samples were taken during the mid-austral summer period, phytoplankton dynamics and nutrient use vary considerably in this region of the Southern Ocean (Whitehouse et al. 1996a, b). Our summer measurements span 51 days between 16 December and 4 February, and there would appear to be some evidence of a seasonal progression. For instance, there was a temperature increase of ~1 C between the two transects measured 1 month apart during cruise JR06 (Whitehouse et al. 1996b; Trathan et al. 1997), and the measurements made during the second transect (February) were the warmest of any measured during the present study. Also, chlorophyll a levels in the AAZ, other than those above the South Georgia shelf, tended to be higher in the early summer. However, there is little consistency between the early January measurements (JR06 T02 and JR11), illustrating the extent of interannual variability. There was spatial variability between the three zones sampled, the stations sampled in each zone, and the various depths sampled at each station. We used a General Linear Model of the data to quantify the magnitude of variability over the entire transect during cruises JR11, 17 and 28, and with additional data from the two transects measured during cruise JR06, a more thorough comparison of the PFZ and AAZ was also considered. Overall, station to station variability within zones was low, especially for temperature and salinity, which implies that the denition of the zones' boundaries was good. The UML was generally about 50 m thick, and all measured parameters were susceptible to substantial change towards the bottom of the UML. Therefore,

greater variability due to depth was occasionally found when the UML was less than 50 m thick. Intercruise variability was by far the greatest factor governing the distribution of near-surface chlorophyll a values (Table 2), and within-cruise dierences were mainly due to sample depth (i.e. between 0 and 50 m). Although the AAZ was usually the zone of highest chlorophyll a concentrations (Fig. 2), considerable overlap between measurements in dierent zones negated zonal importance as a dominating factor. For instance, during cruise JR11, median values were 0.6 and 1.6 mg m)3 in the NZ and the AAZ, respectively, whereas during cruise JR17, values of 1.2 and 9.8 mg m)3 were recorded for the same zones. Interzonal variability was more evident when the comparison was conned to the PFZ and AAZ, but still ranked third below cruise and depth factors (Table 2). Although horizontal phaeopigment distribution varied in a similar way to chlorophyll a, interzonal variability was negligible in both comparisons. Intercruise and depth factors dominated and were probably related to the magnitude of the phytoplankton bloom, and the sub-surface concentration peaks of dead, sinking cells. The zones were dened by their physical attributes; therefore, unsurprisingly, near-surface temperature and salinity measurements were dominated by interzonal dierences (Table 2). This was especially so for temperature, which showed a great deal of intra-zone consistency throughout the ve transect surveys. This zonal dierence persisted through the mid-water depths, especially between 51 and 200 m where >65% of variability over the entire transect length was due to zone, but was less distinctive in waters below 200 m. Interzonal variability was a major factor in silicate and nitrite distribution (Table 2). For silicate, the three zone comparison's high interzonal component was mainly a reection of the particularly low values measured in the NZ. This was less evident in the PFZ-AAZ comparison where values were more alike. However, as with temperature, the zonal dierence persisted through the mid-water depths, where between 51 and 200 m ~65% of variability over the entire transect length was due to zone; again this became less distinctive below 200 m. Depth was also an important variability

Table 2 A comparison of near-surface (050 m) variance components using an adjusted sum of squares (plus ratio expressed as %), in all zones, with cruises JR06 and JR25 excluded, and in the Polar Frontal Zone and Antarctic Zone, with cruise JR25 excluded Chlorophyll a Phaeopigment Temperature Salinity All zones with cruises JR06 and Zone 1.1658 (9) 0.0000 Cruise 8.5526 (65) 0.0828 Station 0.5785 (4) 0.0315 Depth 2.8678 (22) 0.0960 Polar Frontal Zone and Zone 3.7835 (23) Cruise 7.4555 (46) Station 0.5488 (3) Depth 4.6450 (28) JR25 excluded (0) 7.5546 (81) (39) 0.7350 (8) (15) 0.2554 (3) (46) 0.7592 (8) Si(OH)4-Si PO4-P NO3-N NH4-N NO2-N

0.0421 (60) 42.2155 (59) 0.0191 (27) 0.9177 (1) 0.0016 (2) 10.0867 (14) 0.0076 (11) 18.6506 (26)

0.0064 (7) 3.2976 (22) 0.0648 (68) 7.6335 (50) 0.0044 (5) 0.1652 (1) 0.0188 (20) 4.1883 (27)

0.0132 (11) 0.0034 (58) 0.0087 (8) 0.0015 (26) 0.0262 (23) 0.0001 (2) 0.0663 (58) 0.0008 (14) 0.0029 (66) 0.0009 (21) 0.0001 (2) 0.0005 (11)

Antarctic Zone 0.0083 (4) 0.0809 (33) 0.0443 (18) 0.1098 (45)

with cruise JR25 excluded 2.7848 (76) 0.0037 (47) 16.7980 (32) 0.0000 (0) 0.4279 (12) 0.0021 (27) 8.6780 (16) 0.0429 (68) 0.1114 (3) 0.0005 (6) 7.4550 (14) 0.0031 (5) 0.3385 (9) 0.0016 (20) 20.1420 (38) 0.0175 (27)

0.0000 (0) 0.0127 (13) 5.2920 (43) 0.0157 (15) 1.6930 (14) 0.0139 (14) 5.3570 (43) 0.0594 (58)

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component as there is a steep silicate concentration gradient near the bottom of the UML. Also, interstation variability within zone was relatively high, which may reect incursions of other water masses (see below), as well as dierential use by the phytoplankton population (i.e. diatom-dominated or not). Although interzonal variability dominated nitrite distribution, intercruise dierences were also an important factor and were possibly related to the distribution of nitrate and its use by phytoplankton. Although NZ phosphate and nitrate values were generally lower than in the other two zones, interzone variability played a minor role in their distribution. Intercruise variability was the dominant factor, particularly for phosphate, and was related to chlorophyll a distribution and was probably due to phytoplankton use. There were no interzonal dierences evident in the PFZ-AAZ comparison. Ammonium distribution was dominated by depth, which is doubtless due to the sub-surface concentration peaks found at the bottom of the UML. In the mid-water depths around the pycnocline (51200 m), >75% of ammonium concentration variability was due to depth. However, interstation variability was also an important factor and doubtless was a reection of mesoscale dierences in its production and uptake (Fig. 2). Apparent summer nutrient depletion in the surface (030 m) waters A comparison of apparent spring-summer nutrient decits over the scale of our Maurice Ewing Bank transect is of limited quantitative value due to the numerous unmeasured lateral and vertical uxes, such as shifts in frontal system positions, upwellings, sedimentation and preferential nutrient use. For instance, incursions of high-nutrient water in the northernmost stations resulted in them frequently having higher concentrations in summer compared with our ``pre-bloom'' measurements (Fig. 5). However, such a comparison does present an opportunity to consider relative nutrient dynamics through the zones we have sampled. Silicate usage clearly follows the latitudinal gradient of availability, with practically no depletion evident to the north of the transect, but up to ~5 mmol m)3 at station 5 (Fig. 5). The apparent depletion then increased up to ~10 mmol m)3 in the southern PFZ, and ~16 mmol m)3 in the northern AAZ. The greatest depletion in the AAZ appeared to be between stations 17 and 19 (>20 mmol m)3), whereas the least depletion appeared to be nearer the island between stations 20 and 22 (~13 mmol m)3 overall). Therefore given the pre-bloom availability of silicate (Fig. 4), there appeared to be a major surplus in the waters near to South Georgia. Nitrate use appeared to be distinct in ve general regions (Fig. 5). At the northernmost three stations, summer depletion was generally low, and indeed this area may well benet from summer nitrate enhancement due to incursions of nutrient-rich water. Throughout

Fig. 5 The dierence between spring surface (~6 m) and summer near-surface (030 m) silicate, nitrate and phosphate concentrations, along with median dierences (line)

most of the PFZ (stations 512), there appeared to be a depletion of ~8 mmol m)3, and then considerably less in the northernmost part of the AAZ (~4.6 mmol m)3). Highest nitrate use was between stations 18 and 21 (~11 mmol m)3), while nitrate appeared to be underutilised at station 22 above the island shelf (6.2 mmol m)3). Phosphate depletion appeared to be more uniform than both silicate and nitrate depletions (Fig. 5). Stations 14 were subject to minor summer depletions, or repletions due to incursions of highernutrient waters, and then there was a fairly consistent usage to the south. There was a dip in apparent depletions near the fronts at stations 6 and 13 (0.38 and 0.37 mmol m)3 respectively), and again at station 22 over the South Georgia shelf (0.40 mmol m)3). Summer concentrations were reduced by ~0.48 mmol m)3 in the PFZ and ~0.53 mmol m)3 in the AAZ. Overall production observations Clearly phytoplankton production varied considerably over several scales along the Maurice Ewing Bank transect. Chlorophyll a concentrations typical of the open Southern Ocean (Treguer and Jacques 1992) were found during cruise JR11, while atypically high concentrations reminiscent of neritic areas (Holm-Hansen and Mitchell 1991) were found along the transect during cruise JR17. A similar disparity was found along this transect in a previous study when measurements 1 month apart revealed a similar magnitude of dierence (Whitehouse et al. 1996b). Chlorophyll a concentrations in the NZ were typically low, as in much of the Southern Ocean, despite warm, fairly nutrient-rich conditions (Fig. 2). However, in the PFZ and AAZ chlorophyll a concentrations could be considerably higher, and nutrient availability and turnover appeared far greater. The presence of high AAZ open-ocean chlorophyll a

381

concentrations was most likely linked to the hydrographic inuences of South Georgia and the Scotia Ridge; such levels have not previously been recorded at other deep-water Southern Ocean sites, even in the vicinity of frontal systems where enhanced production might be expected. There was some evidence for intraseasonal warming in these surface waters. However, the highest chlorophyll a concentrations early in the season occurred in some of the coolest waters. Apparent nitrate and phosphate depletions were generally constant throughout much of the PFZ, and median spring-summer reductions (7.95 and 0.48 mmol m)3 respectively, 16.6:1) were consistent with Redeld ratio. There was a clear northto-south gradient in silicate use throughout the length of the transect, and in the PFZ its apparent depletion, compared with nitrate and phosphate, was lower than might be anticipated in true Antarctic waters (Priddle et al. 1995). We consider this, and the greater silicate depletions measured in the AAZ, to be attributable to the distribution and relative abundance of diatom species in the phytoplankton community. Silicate depletion was highest in the AAZ, although clearly less so at the southernmost three stations. The highest silicate depletions occurred over deep oceanic water in the AAZ, appeared to coincide with the highest early season (mid-December to early January) chlorophyll a concentrations, and were in or above water potentially inuenced by the Southern ACC Front and WeddellScotia Conuence waters (Orsi et al. 1995). The relationship between silicate, nitrate and phosphate was more complex in the AAZ, where summer ammonium and nitrite concentrations were higher than to the north. Furthermore, the stations over the South Georgia shelf, potentially in a dierent water type (Brandon et al., in press), had higher chlorophyll a concentrations later in the season compared with the stations over deep water. Although some of the highest silicate depletions coincided with high ambient ammonium concentrations, no obvious under-utilisation of nitrate was evident at these stations. However, the highest chlorophyll a concentrations recorded (during cruise JR17) did coincide with high near-surface ammonium concentrations (Fig. 2), along with high sub-surface (50150 m) levels of >1.25 mmol m)3, which would be likely to aect phytoplankton nitrogen preference (see Dortch 1990; Whitehouse et al. 1999). Conclusions There was a consistency in the magnitude of phytoplankton production within the various zones to the north of South Georgia. Chlorophyll a concentrations were far higher in the PFZ than in the waters of the SAZ, but production in the AAZ could be exceptionally high by comparison. However, there appeared to be some sort of discontinuity in the timing of phytoplankton production and nutrient use between the AAZ deep-

water stations and those above the South Georgia shelf. This may be a reection of the changes in relative abundance of dierent phytoplankton species. As already mentioned, chlorophyll a concentrations between 1113 mg m)3 have previously been recorded over deep water to the north of South Georgia (Fryxell et al. 1979; El-Sayed and Weber 1982). Fryxell et al. (1979) speculated that the elevated phytoplankton levels may have been due to seeding from southeast of South Georgia. Our observations would suggest that high chlorophyll a levels are frequently, but not consistently, found over deep water to the north of South Georgia. Although we have no data concerning the likelihood of the area being seeded from the southeast, other observations pointed to the potential impact of WeddellScotia Conuence waters from that area. The AAZ portion of our transect skirted the boundary between ACC water owing from the west and high-silicate waters inuenced by the Weddell-Scotia Conuence, originating from the Antarctic Peninsula, but locally approaching from the east. Silicate is the only major nutrient recorded at growth-limiting concentrations in South Georgia waters (Whitehouse et al. 1996a), and the elevated chlorophyll a levels in the o-shelf waters to the north of the island coincided with high silicate depletions. Therefore it is possible that high-silicate Weddell-Scotia Conuence waters replenish locally depleted waters and maintain large diatom blooms. In addition, Brandon et al. (1999) documented a degree of communication between these o-shelf waters and those on the South Georgia shelf. The shelf waters are rich in ammonium (Whitehouse et al. 1999) and are a likely source of trace elements such as iron, which would further enhance the phytoplankton growth potential in this westerly owing water as and where it meets the warmer, easterly owing waters of the ACC.
Acknowledgements We thank everyone involved in the collection of these data, but especially Alistair Murray for help with the chlorophyll a measurements, and Andy Rees for the cruise JR28 nutrient data. Alistair Murray also advised on the statistical analyses of the data. We are also grateful to two anonymous referees for their helpful comments on the manuscript.

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