Pilot Trophic Model for Subantarctic Water Over the Southern Plateau, New Zealand: a Low Biomass,...

Pilot trophic model for subantarctic water over the

Southern Plateau, New Zealand: a low biomass,

high transfer efficiency system

Janet M. Bradford-Grievea,*, P. Keith Probertb, Scott D. Noddera,David Thompsona, Julie Hallc, Stuart Hanchetd, Philip Boyde,

John Zeldisf, Allan N. Bakerf, Hugh A. Bestg, Niall Broekhuizenc,Simon Childerhousef, Malcolm Clarka, Mark Hadfielda,

Karl Safic, Ian Wilkinsong

aNational Institute of Water and Atmospheric Research, P.O. Box 14901, Kilbirnie, Wellington, New ZealandbDepartment of Marine Science, University of Otago, P.O. Box 56, Dunedin, New Zealand

cNIWA Hamilton, P.O. Box 11115, Hamilton, New ZealanddNIWA Nelson, P.O. Box 893, Nelson, New Zealand

eNIWA Centre for Chemical and Physical Oceanography, Chemistry Department, University of Otago,

P.O. Box 56, Dunedin, New ZealandfNIWA Christchurch, P.O. Box 8602, Christchurch, New Zealand

gDepartment of Conservation, P.O. Box 10420, Wellington, New Zealand

Received 21 June 2002; received in revised form 9 January 2003; accepted 22 January 2003

Abstract

The Southern Plateau subantarctic region, southeast of New Zealand, is an important feeding

area for birds, seals and fish, and a fishing ground for commercially significant species. The

Southern Plateau is a major morphometric feature, covering approximately 433,620 km2 with

average depth of 615 m. The region is noted for its relatively low levels of phytoplankton biomass

and primary production that is iron-limited. In order to evaluate the implications of these attributes

for the functioning of this ecosystem a steady-state, 19-compartment model was constructed using

Ecopath with Ecosim software of Christensen et al. [www.ecopath.org]. The system is driven by

primary production that is primarily governed by the supply of iron and light. The total system

biomass of 6.28 g C m� 2 is very low compared with systems so far modelled with a total system

throughput of 1136 g C m� 2 year� 1. In the model, the Southern Plateau retains 69% of the

biomass in the pelagic system and 99% of total production. Although fish are caught demersally,

0022-0981/03/$ - see front matter D 2003 Elsevier Science B.V. All rights reserved.

doi:10.1016/S0022-0981(03)00045-5

* Corresponding author. Tel.: +64-4-386-0300; fax: +64-4-386-2153.

E-mail address: [email protected] (J.M. Bradford-Grieve).

www.elsevier.com/locate/jembe

Journal of Experimental Marine Biology and Ecology

289 (2003) 223–262

http:\\www.ecopath.org

most of their food is part of production in the pelagic system. Top predators represent about 0.3%

of total biomass and account for about 0.24 g C m� 2 year� 1 of food consumed made up of birds

0.058 g C m� 2 year� 1, seals 0.041 g C m� 2 year� 1, and toothed 0.094 g C m� 2 year� 1 and

baleen whales 0.051 g C m� 2 year� 1. This amounts to 105,803 tonnes carbon over the whole of

the Southern Plateau and is about 17% of the total amount of food eaten by non-mesopelagic fish.

Mean transfer efficiencies between trophic levels II and IV of 23% are at the high end of the range

reported in the literature. In the model, adult fish production is almost completely accounted for by

the fisheries take (32%), consumption by seals (7%), toothed whales (21%), other adult fish

(13%), and squid (20%). Fish and squid catches are at the trophic levels of 4.8 and 5.0,

respectively. The gross efficiency of the fishery is 0.018% (catch/primary production). Although

not all data come from direct knowledge of this system, the model reflects its general

characteristics, namely a low primary production system dominated by the microbial loop, low

sedimentation to the seafloor, high transfer efficiencies, a long food web and supporting high-level

predators.

D 2003 Elsevier Science B.V. All rights reserved.

Keywords: Ecosystem structure; Fish production; New Zealand; Subantarctic; Transfer efficiency; Trophic model

1. Introduction

The Southern Plateau subantarctic region, southeast of New Zealand, is an important

feeding area for birds, seals and fish, and a fishing ground for commercially important

species. The Southern Plateau is a major morphometric feature, covering approximately

433,620 km2 and includes the Campbell Plateau, the Bounty Plateau, Pukaki Rise,

Campbell Rise, and the Auckland Shelf (Carter et al., 1997). The Plateau rises above

sea level at the Auckland, Campbell, Bounty, and Antipodes Islands. Water depths range

from 0–500 m on the rises to 1500 m at the plateau edge (Summerhayes, 1967a,b; Krause

and Cullen, 1970).

The Southern Plateau region is noted for its relatively low levels of phytoplankton

biomass (chlorophyll a) observed from satellite measurements (Murphy et al., 2001) and

field observations and low levels of primary production despite there being plenty of

phytoplankton nutrients such as nitrates (Bradford-Grieve et al., 1997). Recent studies

in subantarctic waters show that some of the reasons for these observations are the very

low levels of dissolved iron and the way this interacts with the availability of light and

silicate (Boyd et al., 1999). The most obvious impact of iron-limited growth is the

dominance of the phytoplankton by small forms. Therefore, the food web in this region is

relatively long. The length of the food web implies that only a small fraction of the low

phytoplankton production reaches the highest trophic levels of large carnivores. Never-

theless, the subantarctic region over the Southern Plateau is of special interest to fishers

who catch several species of fish there, and to conservationists because there are large

populations of seabirds and populations of seals and sea lions most of which feed in the

region. Contrary to this expectation of low efficiency, this study shows the system is

highly efficient.

The aim of the present paper is to identify and quantify the elements of the oceanic

food web that support the birds, mammals and fish of the Southern Plateau, and to

J.M. Bradford-Grieve et al. / J. Exp. Mar. Biol. Ecol. 289 (2003) 223–262224

answer questions such as: What are the implications of the ‘‘microbial’’ system for the

large animals, fish, and birds that depend on production from this subantarctic ecosys-

tem? What are the differences between this system and other types of ecosystems? What

are the priorities for information acquisition to obtain a more reliable picture of this

ecosystem?

To model the Bounty/Campbell ecosystem, the Ecopath software was used (Christensen

et al., 2000). Ecopath has two elements, one that describes the production term and another

for the energy balance for each group. Ecopath bases the model on an assumption of mass

balance over an arbitrary period, usually a year.

2. Materials and methods

2.1. The study area

The area of the Southern Plateau was measured in plan view using digital bathymetric

data and a Geographic Information System to produce closed polygons representing the

various isobaths. The areas between each isobath were calculated and summed. This

plateau is considered to be the area contained by the 1000-m isobath, excluding the

islands, excluding the South Island at the base of the continental slope, and including the

Pukaki Saddle shallower than about 1500 m (Fig. 1). This area is 433,620 km2 of which

9% is 0–250 m, 18% is 250–500 m, 41% is 500–750 m, and 32% is 750–1000 m and

with an average depth of about 615 m.

Salinities and temperatures are nearly uniform over the Campbell Plateau and have

values characteristic of Subantarctic Mode Water in the Southwest Pacific Basin (34.35–

34.40 and 7 jC, respectively) (Morris et al., 2001). Morris et al. (2001) show that winter

surface mixed layers are 200–300 m deep and that there is weak thermal stratification over

the top 500 m across the central plateau region. The Campbell Plateau is a relatively

quiescent region and is bounded almost on all sides by major ocean fronts. Northwest of

the plateau, subantarctic water is bounded by the Southland Front which is part of the

Subtropical Front (e.g. Heath, 1981) which passes northeastward through the Snares

Trough separating the Campbell Plateau from the Snares Shelf immediately south of New

Zealand. Around the southern and eastern flank of the Campbell Plateau, subantarctic

water is separated from less saline Circumpolar flows in the Southwest Pacific Basin by

the Subantarctic Front (Fig. 1). Water flows generally from west to east around the plateau

and follows the bathymetry into cyclonic flow around the Bounty Trough; there is

evidence of weak flows on the plateau, among them an anticyclonic flow around the

Pukaki Rise (Morris et al., 2001).

Macronutrients in subantarctic water over the Campbell Plateau are in good supply

(Levitus et al., 1993; Bradford-Grieve et al., 1997) although NO3 is in excess relative to

DRSi (Zentara and Kamykowski, 1981). Low dissolved iron concentrations are observed

in upper subantarctic water south of 45jS southeast of Tasmania and southeast of New

Zealand (Sedwick et al., 1997; Boyd et al., 1999). Recent phytoplankton physiological

experiments report iron stress as well as light limitation in spring phytoplankton

populations (Boyd et al., 1999). It appears that the low micronutrient status of this region

J.M. Bradford-Grieve et al. / J. Exp. Mar. Biol. Ecol. 289 (2003) 223–262 225

ensures that the ecosystem behaves like a low-nutrient system despite an excess of nitrate

(Bradford-Grieve et al., 1999). Thus the food web is characterised by low chlorophyll

concentrations and small phytoplankton cells. In addition, there is low seasonality in

chlorophyll concentrations, but higher seasonality in phytoplankton and microzooplankton

carbon biomass (Bradford-Grieve et al., 1999). Bradford-Grieve et al. (1999) also report a

large contribution to total community biomass by bacteria and winter bacterial production

to be greater than primary production in the surface 100 m.

In terms of the food web classification of Legendre and Rassoulzadegan (1995), it

appears that, in winter, a ‘‘microbial’’ web or even ‘‘microbial loop’’ system existed

because microbial rather than autotrophic processes dominated (Bradford-Grieve et al.,

1999). The spring food web is classified as ‘‘microbial’’ as phytoplankton made a larger

contribution to the creation of living organic particles than in winter, but material was still

being directed to small heterotrophs such that microzooplankton biomass was greater than

that of mesozooplankton. There is a very low sedimentation of particles from this

‘‘microbial’’ system (Nodder and Northcote, 2001; H. Neil, personal communication).

Fig. 1. Map of the region showing the Southern Plateau (Bounty/Campbell Plateau) and the water masses and

fronts in relation to the Plateau.


2.2. Modelling approach

The modelling approach of Ecopath with Ecosim is given by Christensen et al. (2000).

In this model, a system of simultaneous linear equations can be solved using standard

matrix algebra:

B1PB1EE1 � B1QB1DC11 � B2QB2DC21 � . . .� BnQBnDCn1 � EX1 ¼ 0

B2PB2EE2 � B1QB1DC12 � B2QB2DC22 � . . .� BnQBnDCn2 � EX2 ¼ 0

. . .

BnPBnEEn � B1QB1DC1n � B2QB2DC2n � . . .� BnQBnDCjn � EXn ¼ 0

where Bn is the biomass of element (compartment or box) n, PBn is the production/

biomass ratio, QBn is the consumption/biomass ratio, DCjn is the fraction of prey (n) in

the average diet of predator j, EEn is the ecotrophic efficiency of n, and EXn is the export

of n.

Ecopath sets up a system with as many linear equations as there are groups in the

system and it solves the set for one of the following parameters for each group: biomass

(B); production/biomass ratio (P/B); consumption/biomass ratio (Q/B); ecotrophic effi-

ciency (EE) = fraction of production that is used in the system, i.e. either passed up the

food web, used for biomass accumulation, migration or export. As well as three of the

above parameters, the following parameters must also be entered for all groups: catches

(and discards); net migration rate (here set to 0); biomass accumulation rate (here set to 0);

assimilation rate; diet compositions; fate of detritus. Energy balance within a compartment

is ensured using the equation: consumption = production + respiration + unassimilated

food.

2.3. Model groups (compartments)

From our knowledge of the functioning of the early food web and the diet of high-level

predators, we define the food web of the Bounty Campbell Plateau as having the following

functional compartments. All available information on biomass, catch, P/B ratios, and

consumption rates (Q/B) have been assembled from our own research data, the literature,

and catch statistics. Then species of similar sizes, diets, and consumption rates, etc., were

aggregated within a compartment (Fig. 2). The resulting 19-compartment model consists

of (1) birds, (2) seals and sea lions (3) toothed whales, (4) fish (adult), (5) fish (juvenile)

(6) baleen whales, (7) squid, (8) mesopelagic fish, (9) macrozooplankton, (10) macro-

benthos, (11) mesozooplankton, (12) ciliates, (13) heterotrophic flagellates, (14) meio-

benthos, (15) bacteria (water column), (16) bacteria (sediment), (17) phytoplankton, (18)

detritus water (column), and (19) detritus (benthic).

Biomasses that were not already available in terms of carbon were converted to carbon

using the following relationships. Birds and mammals: 10% of wet weight is assumed


to be carbon; fish and squid: 10% of wet weight is carbon assuming that carbon

content is about 50% of dry weight and that dry weight is about 20% of wet weight

(Vinogradov, 1953); macro- and mesozooplankton: log(WW) =� 1.537 + 0.852log(C)

Fig. 2. Trophic model for the Southern Plateau, New Zealand. The box size is proportional to the square root of

the compartment biomass. All flows are in g C m� 2.


and log(DW) = 0.499 + 0.991log(C), where WW=wet weight, DW= dry weight, and

C = carbon (Weibe, 1988).

2.4. Input data

The origin of input data is given in Appendix A and is summarised in Tables 1–3.

2.5. Balancing the model

Initially ecotrophic efficiencies (the fraction of total production consumed by predators

or exported from the system) of a number of early food web compartments were >1. Part

Table 1

Prey–predator matrix used in model

Prey/predator 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16

(1) Birds

(2) Seals, etc.

(3) Toothed whales

(4) Fish (adult) 0.25 0.3 0.015 0.06

(5) Fish (juvenile) 0.05 0.040 0.07

(6) Baleen whales

(7) Squid 0.30 0.30 0.7 0.015 0.10

(8) Mesopel. fish 0.31 0.45 0.155 0.40 0.07

(9) Macrozoopl. 0.34 0.615 0.40 0.50 0.70 0.38 0.05

(10) Macrobenthos 0.160 0.05

(11) Mesozoopl. 0.60 0.10 0.62 0.55 0.15

(12) Ciliates 0.20

(13) Het. flagellates 0.15 0.55 0.60 0.10

(14) Meiobenthos 0.30 0.20

(15) Bacteria (col.) 0.20

(16) Bacteria (sed.) 0.60 0.80

(17) Phytoplankton 0.30 0.10 0.40 0.70

(18) Detritus (col.) 1.0

(19) Detritus (sed.) 1.0

Sum 1.00 1.00 1.0 1.00 1.00 1.00 1.00 1.0 1.00 1.00 1.00 1.00 1.00 1.0 1.0

Numbers represent biomass fractions of food ingested.

Table 2

Average annual catch (wet tonnes) by species from stocks inhabiting the Campbell Plateau (data from Annala

et al., 1999)

Species Range Average N

Hake 1803–3956 2999 (area 1) 9

Hoki 10,000–34,000 subantarctic 19,300 subantarctic 9

76,000–165,000 west coast SI 109,667 west coast SI 9

Ling 935–7510 4946 (area 6) 9

Orange roughy 200–3850 1158 6

Oreos 2528–5645 4457 4

Southern blue whiting 13.7–76.2 30,117 9

N= number of years averaged. SI = South Island.


Table 3

Input parameters and calculated parameters (in bold) for the Southern Plateau

Catch Trophic

level

Biomass P/B Q/B EE P/Q Net

efficiency

R A R/A P/R R/B Flow to

detritus

(1) Birds 0.00000105 5.3 0.00158 0.30 36.5 0.002 0.008 0.010 0.046 0.046 0.990 0.010 28.900 0.012

(2) Seals 0.00000066 5.7 0.0009 0.22 46.0 0.04 0.005 0.006 0.033 0.033 0.994 0.006 36.580 0.008

(3) Toothed whales 5.9 0.0085 0.06 11.0 0.00 0.005 0.007 0.074 0.075 0.993 0.007 8.740 0.019

(4) Fish (adult) 0.0153 4.8 0.4517 0.30 2.60 0.935 0.115 0.141 0.805 0.940 0.856 0.169 1.760 0.244

(5) Fish (juvenile) 4.5 0.086 1.00 3.00 0.945 0.333 0.417 0.120 0.206 0.583 0.714 1.400 0.056

(6) Baleen whales 5.0 0.004 0.038 12.8 0.00 0.003 0.004 0.041 0.041 0.996 0.004 10.202 0.010

(7) Squid 0.0042 5.0 0.0204 8.00 22.0 0.992 0.364 0.455 0.196 0.359 0.545 0.833 9.600 0.091

(8) Mesopelagic fish 4.5 0.285 1.00 16.0 0.95 0.063 0.104 2.449 2.734 0.896 0.116 8.600 1.837

(9) Macrozooplankton 3.6 0.311 10.0 33.0 0.95 0.303 0.505 3.046 6.154 0.495 1.020 9.800 4.258

(10) Macrobenthos 3.5 0.25 1.00 2.86 0.895 0.350 0.583 0.179 0.429 0.417 1.400 0.714 0.312

(11) Mesozooplankton 3.5 1.058 20.00 57.14 0.836 0.350 0.538 18.137 39.297 0.462 1.167 17.143 24.625

(12) Ciliates 2.8 0.167 88.00 247.89 0.927 0.355 0.507 14.282 28.978 0.493 1.029 85.521 13.485

(13) Heterotr. flagellates 2.3 0.307 292 829.55 0.932 0.352 0.503 88.625 178.269 0.497 1.011 288.682 82.288

(14) Meiobenthos 3.3 0.1 10.00 32.26 0.859 0.310 0.443 1.258 2.258 0.557 0.795 12.581 1.108

(15) Bacteria (column) 2.0 0.598 87.40 383.33 0.975 0.228 0.289 128.829 183.094 0.711 0.406 215.433 49.470

(16) Bacteria (sediment) 2.0 1.5 2.508 8.36 0.80 0.300 0.375 6.270 10.032 0.625 0.600 4.180 3.261

(17) Phytoplankton 1.0 1.253 248 0.769 61.409

(18) Detritus (column) 1.0 572 0.963 8.781

(19) Detritus (sediment) 1.0 433 0.931 0.000

Catch, biomass (B), production (P), respiration (R), assimilation (A), and flow to detritus in terms of g C m� 2 year� 1. P/B, Q/B, R/B year� 1. EE, P/Q, net efficiency

(P/A), R/A, and P/R are dimensionless.

J.M.Bradford-G

rieveet

al./J.

Exp.Mar.Biol.Ecol.289(2003)223–262

230

of this problem appeared to be that phytoplankton production relative to biomass (P/B), as

calculated from a limited set of original data, was not great enough to support the biomass

that immediately depended on it, as defined in the model. Therefore we chose a value for

primary production that was defined in proportion to bacterial production based on the

conclusions of Ducklow (2000).

Detritus derived from intertidal primary producers (macroalgae) was not taken into

account in the model although it will be important locally around the islands on the

Southern Plateau. Macroalgae can make a direct input to the deep sediment (Reichardt,

1987) but of most interest is the production of fine particulate detrital material and DOC

into the water column (Mazure and Field, 1980 and references therein). Kelp beds may be

highly productive with their productivity per unit area often being greater than the

phytoplankton by a factor of 10 (Mann, 1973). The particles remain suspended in the

water for a time and are acted on by bacteria (Laycock, 1974). Mazure and Field (1980)

estimate that the total amount of carbon available to suspension-feeders from kelp plants in

the Benguela current kelp beds may be in the order of 1200–1600 g C m� 2 year� 1. It is

likely that subantarctic algae do not produce as much detritus as this, but as yet no data are

available on which to evaluate their contribution to this system.

Initially we did not believe that annual P/B for sediment bacteria was as low as

experimental data suggested and chose a literature-derived value of 11. The model

produced a very low ecotrophic efficiency for bacteria so it was decided to let the model

calculate P/B by assuming an ecotrophic efficiency of 0.8 and P/Q of 0.3. The

subsequently calculated annual P/B for sediment bacteria of 2.5 is nearer the experimen-

tally derived P/B of 1 than the literature value of 11.

Slight changes in the diet of some compartments (Table 1) reduced the ecotrophic

efficiencies of their prey to < 1.

Initially there was not enough material flowing to the detritus compartments. Changes

in the ratio of unassimilated food/food consumption corrected this. Final balancing was

achieved by choosing relatively high production/consumption ratios for ciliates, hetero-

trophic flagellates, and meiobenthos (Table 3).

3. Results and discussion

3.1. Evaluation of data

A summary of all the final input data and the calculated parameters is found in Table 3.

Although we have detailed data available for subantarctic water just north of the

Plateau, we do not know if these data are representative of the Plateau itself. Also, a

number of the parameters were estimated from literature values. The absence of data on

production/biomass ratios and quantitative food consumption in relation to biomass for

commercial fish species on the Plateau was particularly conspicuous. Our knowledge of

annual and interannual variability on the Plateau is very limited therefore our estimation of

annual averages has to be considered approximate.

The primary production figure used in the model is probably overestimated. Boyd et

al. (unpublished manuscript) compare in situ primary productivity measurements and


primary production estimated from SeaWiFS data and the algorithm of Behrenfeld and

Falkowski (1997). This method was modified for New Zealand waters using sea surface

temperature data from AVHRR and local optical spectral attenuation data. This

technique showed a robust relationship between the measured and estimated data and

gave an annual production of 142.5 g C m� 2 year� 1 for a box which takes in most, but

not all, of the Southern Plateau region (Murphy et al., 2001). This means that the

localised high chlorophyll a in the vicinity of the Auckland Islands (>1 mg m� 3) and

the calculated column integrated primary production of 2–2.5 g C m� 2 day� 1 are not

included in the above estimate of average annual primary production for the Southern

Plateau. Thus, although average annual primary production is likely to be greater than

142.5 g C m� 2 year� 1, it is probably not as great as the 265 g C m� 2 year� 1 use in the

model.

The model was balanced by using what may be unrealistically high P/Q ratios for some

compartments (ciliates: 0.355; heterotrophic flagellates: 0.352; macrobenthos: 0.35;

meiobenthos: 0.31). Total daily consumption/biomass for microzooplankton is estimated

from a limited set of dilution experiments at 10 m for microzooplankton grazing on < 200

Am phytoplankton, ciliates grazing on heterotrophic flagellates, and heterotrophic flag-

ellates grazing on bacteria (J.H., unpublished data). These data were converted to an

annual basis and the P/Q ratio determined using a P/B of 220. The P/Q values average

0.242 for ciliates and heterotrophic flagellates combined. This is less than the 0.35 used in

the model for ciliates and heterotrophic flagellates but may be justified on the grounds that

the above experimental estimate is from water at 10 m and probably does not reflect the

average situation down the water column.

Experimental data on the iron-limited system of the North Pacific suggests that this

particular nutrient limitation impacts a number of ecological parameters. P/Q for

heterotrophic bacteria (Tortell et al., 1966) was 0.12–0.36 in iron-deficient conditions

and for heterotrophic marine microflagellates (Chase and Price, 1997) was 0.18. In the

model, P/Q for water column bacteria (0.228) fits within these values and the range of

experimentally derived values, whereas P/Q for microzooplankton is less than the P/Q

used here (0.352).

Total consumption by microzooplankton in the NE Pacific of bacteria is greater than

that on phytoplankton (58% and 26% of the total ration, respectively) (Rivkin et al.,

1999). Data from Hall et al. (1999) and Bradford-Grieve et al. (1997) suggest that bacteria

and picophytoplankton each make up about 50% of the total ration of heterotrophic

flagellates. Nevertheless, the model could be balanced only with 20% of heterotrophic

flagellates diet being made up of bacteria. The reason for this may be located in the

possible underestimation of water column bacterial biomass. It was established that

bacterial P/B and B are more important than the substrate for bacterial growth in

determining the availability of bacteria to heterotrophic flagellates. That is, importing

detritus from macroalgae does not make a difference to the carrying capacity of the

pelagic system as defined.

For some parameters we have direct measurements. For example, the input of carbon

sedimented to the seafloor (1.23 g C m� 2 year� 1) is less than the 8.8 g C m� 2 year� 1

that the model sends to the seafloor to fuel the activities of the bacteria, meiobenthos, and

macrobenthos. This could be a result of overestimating their biomasses through using data


from the south side of Chatham Rise. Further data are needed to decide whether the

current choice of parameter values for the Southern Plateau region is a serious flaw in the

model.

There was a discrepancy between measured annual P/B for sediment bacteria of 1 and

literature values of about 11. The model calculates that P/B for sediment bacteria of 2.5 is

all that is necessary to satisfy the demand of the benthic system, implying that the

measured P/B for sediment bacteria is not as unrealistic as initially thought.

Many of the other parameter estimates appear to be reasonable. The gross efficiencies

(GE or P/Q) invertebrates are between 0.3 and 0.36 are at the high end of the range

reported in the literature (e.g. Parsons et al., 1984 [K1 (as %) in Table 25]). The P/Q for

mesopelagic fish appears to be very low and is the result of the particular choice of P/B

and Q/B for this compartment. The production/respiration ratios (P/R) are more or less

as expected for those groups of organisms where there are data (Humphrey, 1979). In

other groups, P/R seems high at about 1 but this may not be surprising given the

variability in Humphrey’s (1979) data. It is possible that in this model not enough

energy from food consumption is set aside for respiration and that P/B and/or the

proportion of food that is unassimilated are too high in a number of groups. There is

evidence that this tightly coupled ecosystem may not always be able to support all

animals at optimal food consumption and growth. Since we are assuming optimum

growth in selecting production and food consumption parameters, this may be the reason

that in the model there is not enough energy for respiration. Evidence for this is seen in

the data on growth of the 1991 year class of southern blue whiting (Fig. 3). Larvae of

this species survived in particularly large numbers in 1991. Subsequently, they grew

slower than some previous year classes, suggesting that, in this model, lower P/B and Q/

B would have to have been assigned to reflect the true state of affairs. The low

respiration/biomass for mesozooplankton may be partly accounted for by the fact that

one of the large calanoid copepods in this system (Neocalanus tonsus) spends from late

January/February until September below 500 m where it enters a type of diapause

(Ohman, 1987).

The model calculates that there is 0.31 g C m� 2 of macrozooplankton. This is

converted to 0.99 g DW m� 2 using the relationship given by Weibe (1988). This

concentration is within the range of biomass values given by Pakhomov et al. (1994,

1996) for subantarctic open water of 0.01–4.4 g DW m� 2 and around islands of 0.01–

Fig. 3. Weight at age by cohort for 1987 and 1991 year classes of southern blue whiting (S.H., unpublished data).


2.86 g DW m� 2. The biomass of mesopelagic fish, calculated by the model (0.29 g C

m� 2) for the Southern Plateau, is 30% more than the micronekton biomass of 0.22 g C

m� 2 on the mid-slope region off southern Tasmania, Australia (Williams and Koslow,

1997), north of the Subtropical Front.

3.2. Biomass, consumption, and production

Omitting the large predators (birds, seals, whales), the pelagic domain contains 69% of

total living biomass and benthic compartments (including 0.07 g C m� 2 demersally

feeding fish (16% of biomass)) contain 31%. The benthic compartments account for food

intake of 16.67 g C m� 2 year� 1 which is < 3% of total non-detrital consumption.

Heterotrophic flagellates had the greatest food intake (255 g C m� 2 year� 1) followed

by water column bacteria (229 g C m� 2 year� 1), mesozooplankton (60 g C m� 2 year� 1),

and ciliates (41 g C m� 2 year� 1). In our model, bacteria (sediment plus water column)

utilise 52 g C m� 2 year� 1 of detritus of which 94% is derived from water column

processes. Twenty six percent of water column detritus is derived directly from phyto-

plankton.

The fish compartments (adult fish, juvenile fish) represent about 9% of total biomass

and account for about 1.43 g C m� 2 year� 1 of food intake. Fish and squid catches are at

the trophic levels of 4.8 and 5.0, respectively (Table 3).

According to model results, a significant proportion (56%) of primary production and

water column heterotrophic production is not directly used but enters the detritus pool that

is used by water column and sediment bacteria. The Southern Plateau retains 69% of the

biomass in the pelagic system but 99% of total production. Although fish are caught

demersally, most of their food is part of production in the pelagic system.

The small proportion of fish depending on the benthos for food appears to agree with

conclusions of other bathyal habitats. Haedrich and Merrett (1992) determine that only

13% of fish species in the Porcupine Seabight depend on benthos for food, and a number

of other studies show that slope fish species feed on the mesopelagic fauna (Mauchline and

Gordon, 1984, 1986, 1991; Sedberry and Musick, 1978; Marshall, 1979; Gordon and

Mauchline, 1990). Blaber and Bulman (1987) also show that fish of the eastern Tasmanian

slope (420–550 m) depend on mesopelagic fish resources.

Top predators represent about 0.26% of total biomass and account for about 0.24 g C

m� 2 year� 1 of food consumed made up of birds 0.058 g C m� 2 year� 1, seals 0.041 g C

m� 2 year� 1, and toothed 0.094 g C m� 2 year� 1 and baleen whales 0.051 g C m� 2

year� 1. This amounts to 105,803 tonnes carbon over the whole of the Southern Plateau

and is about 17% of the total amount of food (621,399 tonnes of carbon) eaten by fish

(non-mesopelagic).

The fact that fisheries discards have been directed to ‘‘detritus’’ in the model gives an

erroneous picture of the impact of fishing on the food of some sea birds that tend to

congregate around fishing vessels. Nevertheless, of the estimated >10 million sea birds

that live in the area (D.T., unpublished data), only a small fraction benefits from the

presence of fishing vessels. Therefore, the omission of discards from the food of seabirds

is considered to be immaterial to the functioning of the ecosystem as a whole and birds in

particular.


The partitioning of total adult fish production in the model among those compartments

utilising it (Fig. 4) shows that fisheries take the greatest proportion (32%). The remaining

proportions are 7% consumed by seals, 21% by toothed whales, 13% by other adult fish,

20% by squid, with 7% dying of other causes (give the steady state assumptions). These

proportions do not necessarily absolutely represent the Southern Plateau system but give a

picture of what might be occurring. It would be instructive to know the fate of production

of the most important fish species, but this would require the collection of data specific to

New Zealand’s fish species and that biomasses, production, and consumption information

on the major predators would have to be estimated more accurately.

3.3. Transfer efficiency

Ecosystem components can be grouped into discrete trophic levels and transfer

efficiencies estimated (Lindeman, 1942). Transfer efficiency is the fraction of total flows

at each trophic level (throughput) that are either exported or transferred to another trophic

level through consumption. Transfer efficiencies are greater at the beginning of the food

web compared with higher trophic levels because of intrinsic differences in metabolic

characteristics of organisms at different levels in the food web (e.g. R/A and P/R in Table

3). Transfer efficiencies in the Southern Plateau ecosystem (Table 4) are at the high end of

the range reported in the literature (e.g. Parsons et al., 1984). They are higher than the

Table 4

Transfer efficiencies (the proportion of energy transferred from one trophic level to the next) for each trophic level

Source/trophic level I II III IV V VI VII VIII IX

Producer – 25.8 22.8 19.2 14.4 8.3 4.9

Detritus – 22.3 24.5 23.6 19.2 14.9 8.4 5.0

All flows – 23.9 23.7 21.5 17.2 12.6 7.0 4.6 1.5

Proportion of total flow originating from detritus: 0.51.

Transfer efficiencies (calculated as geometric mean for TL II– IV): from primary producers, 22.4%; from detritus,

23.5%; total, 23.0%.

Fig. 4. Partitioning of adult fish production amongst those compartments consuming it.


upper end of average values (8–15%) reported by Christensen and Pauly (1993), Wolff

(1994), and Wolff et al. (1996).

3.4. Mixed trophic impacts

Direct and indirect interactions within the food web can be evaluated by changing the

biomass of a group and assessing the effect this change will have on other groups in the

system assuming the trophic structure remains the same (Fig. 5). The mixed trophic impact

(MTI) (scaled � 1 to + 1) for each group is calculated using the approach in Ecopath with

Ecosim (Christensen et al., 2000) which is based on the work of Ulanowicz and Puccia

(1990).

In the model, a small increase in the biomass of top predators (birds, seals, toothed

whales, adult fish) all have a negative impact on the biomass of their preferred prey, but it

is the adult fish that have the greatest impact on their prey (apart from macrozooplankton)

and also negatively impact birds and seals as they are directly competing for resources. All

compartments have a negative effect on themselves probably reflecting competition for

resources. An increase in phytoplankton biomass positively impacts most other compart-

ments apart from itself and water column bacteria which are the prey of heterotrophic

flagellates whose biomass is increased by the increase in phytoplankton biomass. An

increase in the biomass of mesozooplankton has the greatest number of positive impacts

on other compartments ranging from small to moderate. Of these impacts, it appears that

the increase in macrozooplankton biomass has the greatest (nevertheless small) impact on

adult fish biomass. An increase in adult fish has a large negative impact on birds, seals,

presumably because of competition for food.

The greatest impact is noted on bird and seal biomass if the fishing fleet is increased

and the resulting accidental bycatch and competition of the fisheries for the food of birds

and seals, as portrayed in the model, is increased. It is also noted that an increase in the

fishing fleet has a positive impact on mesopelagic fish and macrobenthos that make up the

food of fish. These changes cannot be interpreted as predictions as the model, as used,

does not accommodate changes in diet composition that would result. Also, if we were to

identify the exact diets of birds, seals, and individual species of fish in the fishery in more

detail, we might find that there is not the degree of competition for the same food that the

model currently implies.

3.5. Summary statistics

The total throughput (the sum of all flows: consumption, exports, respiratory flows, and

flows to detritus) at each trophic level is 1136 g C m� 2 year� 1 (Table 5), which is low and

similar to throughput within the Golfo Dulce (Wolff et al., 1996). About 97% of

throughput is achieved from trophic levels I to III: 50% from levels I to II plus 38%

from levels II–III. About 54% of the total is due to consumption, < 0.1% is exported

(sedimentation and fishery), 22% flows into detritus, and 23% is respired (Fig. 6).

The total primary production/respiration (P/R) ratio calculated by the model is 1.004

(Table 5), which is similar to estimates obtained from in situ primary production and

plankton respiration data (P/R = 0.9–1.2, see Wolff et al., 1996). That is, the Southern


Fig. 5. Mixed trophic impacts (MTIi, j=DCi, j� FCi, j) on the Southern Plateau (where DC is diet composition, FC

is a host composition term giving the proportion of the predation on j that is due to i as predator). Direct and

indirect impacts of a small increase in the biomass of each of the impacting groups (vertical axis) on every other

group (horizontal axis) are shown. Positive impacts are shown above the line, negative below. Impacts are relative

and comparable between groups.


Plateau lies within the range of open ocean systems where productivity and respiration

tends to be balanced (P/R = 1 Christensen and Pauly, 1993) and where primary production

is comparatively low. The system biomass of 6.219 g C m� 2 is very low compared with

the 41 trophic models presented by Christensen and Pauly (1993) and Wolff (1994), and

even lower than in a tropical fjord-like embayment (Wolff et al., 1996).

An ecosystem picture emerges in which energy fluxes, and to a lesser extent biomass, is

concentrated in the pelagic environment. The mean level of fisheries (4.5) is similar to that

found in the tropical Golfo Dulce (Costa Rica) (5.3) and reflects the fact that the Southern

Table 5

Summary statistics for the Southern Plateau ecosystem model

Parameter Value Units Comment

Sum of all consumption 610 g C m� 2 year� 1

Sum of all exports 0.971 g C m� 2 year� 1 sediment and fishery

Sum of all respiration flows 264 g C m� 2 year� 1

Sum of all flows into detritus 251 g C m� 2 year� 1

Total system throughput 1136 g C m� 2 year� 1

Sum of all production 451 g C m� 2 year� 1

Fishery’s mean trophic level 4.48

Its gross efficiency (catch/primary production) 0.000181

Calculated total net primary production 265 g C m� 2 year� 1

Total primary production/total respiration 1.004

Total biomass/total throughput 0.005

Total biomass (excluding detritus) 6.219 g C m� 2

Total catches 0.048 g C m� 2 year� 1

Throughput cycled (excluding detritus) 35.28 g C m� 2 year� 1

Finn’s cycling index 19.75 % of total throughput

Average path length 5.79

Fig. 6. Partitioning of throughput in the system among consumption by predators, exports, flows to detritus, and

respiration.


Plateau fishery targets predatory fish feeding mainly on prey dependant on the pelagic

system, which is itself oligotrophic in nature because of iron limitation and dominated by

microbial processes. The very low gross efficiency of the fishery (catch/primary produc-

tivity) (0.018%) (Table 5) shows most of the system’s production, which occurs in the

pelagic part of the system, is not harvested in this particular part of the New Zealand EEZ

but is going to sustain the system as a whole.

Another feature of this type of system should be kept in mind. Fish production should

become increasingly sensitive to variations in primary production as primary production

decreases (Iverson, 1990). Iverson calculates that F 20% variation in annual phytoplank-

ton production could result in F 24% variation of fish production at 250 g C m� 2 year� 1

primary production, whereas at 50 g C m� 2 year� 1 primary production, there would be a

F 74% variation in fish production. He concludes that ‘‘the effects of interannual

variations in phytoplankton production on fish production should result in more difficult

fisheries management in environments with low phytoplankton production values’’. The

truth of this generalisation needs to be further investigated for the Southern Plateau as this

region plays an important part in the life cycle of hoki. It is possible that the condition of

hoki adults that go to breed off the west coast of South Island may be directly impacted by

variations in primary productivity or competition for food among elements of the

ecosystem over the Plateau and, as noted above, there is recent evidence of food limitation

of the growth of southern blue whiting in this system.

By definition, the Southern Plateau system is high on the ‘‘maturity’’ scale as we have

assumed steady state and there is no biomass accumulation, i.e. P/R = 1.0 (Odum, 1971). A

high mean total transfer efficiency of 23% exceeds that of any of the systems reported by

Christensen and Pauly (1993). Han (1997) considers that total system standing stock

(TSS)/total system throughput (TST) is the best measure of system structure and state. In

the Southern Plateau system, this ratio is 0.006 year� 1 which is even lower than the Golfo

Dulce (Wolff et al., 1996) and suggests this ecosystem, as modelled, is highly complex

compared with other systems that have been modelled (Table 6). The degree of energy

cycling (assumed to increase as systems ‘‘mature’’ (Odum, 1971)) compared with the 41

systems reported by Christensen and Pauly (1993) shows that Southern Plateau is

intermediate to high (Finn’s cycling index (FCI) = 19.75%; average path length = 5.79).

Nevertheless, these conclusions may be partly an artefact of the detail with which the

zooplankton and microbial system has been described in the present model, as other

Ecopath models referred to here, divide the fish compartment but tend to aggregate the

zooplankton and microbial compartments.

3.6. Sensitivity analysis

The sensitivity analysis routine was used to explore the sensitivity of the model to the

input parameters used in the model. Input parameters are varied in the routine independ-

ently in steps of 10% from � 50% to + 50% of the mean estimates used as input to the

model. Altering input parameters of a group often has large effects on the output

parameters of that group. As expected, changing biomass of most groups, usually by

� 20% or + 30%, had a >20% impact on ecotrophic efficiencies (EEs) of the same groups

(except toothed and baleen whales, mesopelagic fish, macrozoopolankton). Changing P/B


Table 6

Ecosystem maturity statistics

Ecosystem TSS

(g C m� 2)

TST

(g C m� 2 year� 1)

TSS/TST

(year� 1)

FCI

(%)

Average

path

length

Fishery

mean

trophic

level

Gross

efficiency

fishery

(%)

Average

transfer

efficiency

levels

II– IV

Reference

Southern Plateau, NZ 6.22 1136 0.006 19.8 5.79 4.84 0.02 23.0 This study

Golfo Dulce, Costa Rica 10.43 1405 0.007 18.9 3.37 Wolff et al., 1996

Tongoy Bay, Chile 236.3 20,835 0.011 10.1 4.91 3.6 0.89 13.0 Wolff, 1994

Venuzuela shelf 122.10 7621 0.016 1.6 – – – – Mendoza, 1993

Weddell Sea, Antarctica 16.9 261 0.065 18.8 3.5 – – – Jarre-Teichmann et al., 1997

South Benguela, 1989 36.0 2332.7 0.015 – 3.19 3.48 – 10.6 Jarre-Teichmann et al., 1998

South Benguela, 1995–1999 56.6 2081.8 0.027 – 2.78 3.66 0.04 27.4 Jarre-Teichmann et al., 1998

TSS= total system standing stock (biomass); TST= total system throughput; FCI = Finn’s cycling index; gross efficiency of fishery = catch/primary production.

J.M.Bradford-G

rieveet

al./J.

Exp.Mar.Biol.Ecol.289(2003)223–262

240

of most groups by � 20% or + 30% had a >20% effect on the EEs of the same groups

(except for toothed and baleen whales).

The largest impacts of one group on another are seen when relatively low percentage

changes (F 20%) in input parameters of adult and juvenile fish, squid, mesopelagic fish,

macrozooplankton, macrobenthos, ciliates, heterotrophic flagellates, and bacteria are

made. Among the groups impacted by >20%, apart from themselves, are mesopelagic

fish, macrobenthos, macrozooplankton, ciliates, and phytoplankton which are the most

sensitive groups that are impacted by a 20% or 30% change in specific parameters of the

impacting group.

3.7. Limitations and strengths of the approach

How representative of the Southern Plateau ecosystem is this model? At best it can be

seen as preliminary as a number of the parameters were estimated from literature values or

were derived from data collected north of the study region. The fact that the model could

be balanced only by using relatively high P/Q ratios may be a true characteristic of the

subantarctic system. For example, Ohman and Runge (1994) hinted that a calanoid

copepod (Calanus finmarchicus) may need to consume less food when feeding on

microzooplankon compared with the same species feeding on phytoplankton.

All of the gaps in our knowledge of the Southern Plateau ecosystem, identified in the

discussion and in Appendix A, suggest priorities for further work to quantify these

parameters for the Southern Plateau region especially in the area of obtaining production,

food, and respiration characteristics for New Zealand’s major fisheries species.

The assumption of steady state for this system may have caused some of the difficulties

in balancing this model. There is interannual variability in surface chlorophyll a in the

Southern Plateau region (Murphy et al., 2001). Therefore, there is likely to be similar

interannual variability in primary production, perhaps evidenced by variable recruitment to

the southern blue whiting fishery (Hanchet et al., 1998) and less than optimal growth of

the 1991 year class of southern blue whiting that survived in particularly large numbers

(Fig. 3). Because the data used here have been collected in different years, we may be

getting only a general picture.

Nevertheless, we are confident the model reflects the general characteristics of this

system. That is, being a low primary production system dominated by the microbial loop

and with low sedimentation to the seafloor, it is a system with high transfer efficiencies

that supports a range of trophic levels and high-level predators.

Acknowledgements

This study was funded by the Foundation for Research Science and Technology, New

Zealand, Contract No. CO1214. We thank Dr. Helen Neil, Dr. Rosie Hurst, and Marieke

van Kooten, all of NIWA, and Dr. W. Balzer, University of Bremen, Germany, for

providing as yet unpublished data that is acknowledged in the text. The authors are also

grateful to Dr. Lynne Shannon, of Marine and Coastal Management, South Africa, for

sharing her knowledge and experience of this type of modelling. [RW]


Appendix A. Input data

A.1. Birds

Biomass data were collated on the populations of the 11 most common seabird

species (Diomedea antipodensis, Diomedea gibsoni, Diomedea epomophora, Thalas-

sarche impavida, Thalassarche steadi, Thalassarche salvini, Puffinus griseus, Procel-

laria aequinoctalis, Pachyptila desolata, Pterodroma lessonii, Eudyptes sclateri) that

breed on the Bounty/Campbell Plateau (Robertson and Bell, 1984; Gales, 1998; Taylor,

2000), their wet body weights (Marchant and Higgins, 1990), months of residence in the

area (Marchant and Higgins, 1990), and breeding success (data cited in Marchant and

Higgins, 1990). It is assumed that the minor species make a negligible contribution to

seabird biomass. It is also assumed that the populations of smaller sea birds breeding in

the Bounty/Campbell Plateau region also feed there. We do not have the data to decide

how reasonable these assumptions are. It is known that some large species feed outside

the region (e.g. Walker, 1995; Waugh et al., 2000). We assume that 10% of wet weight

is carbon, based on the data for fish (Vinogradov, 1953). Thus the mean annual biomass

of seabirds is assumed to be 0.00158 g C m� 2. Production to biomass ratios were

calculated by using available data on chick maximum weight, number of months to

attain this weight, breeding success, number of breeding pairs, and total population

biomass. An upper bound to annual P/B was calculated from: [(no. of breeding

pairs� no. of eggs laid�maximum chick weight)/total biomass of population]/(no.

of months to maximum weight if over 12 months/12). A lower bound to annual P/B

was calculated from: [(no. of breeding pairs� no. of eggs laid� breeding success�maximum chick weight)/total biomass of population]/(no. of months to maximum

weight if over 12 months/12). A P/B of 0.30 year� 1 is used although Crawford et al.

(1991) use 0.2 year� 1 for southwest African seabirds and Wolff (1994) uses 0.07 year� 1

for northern Chile seabirds.

Annual consumption to biomass ratios were calculated from available data on food

requirements in the literature (e.g. Woehler and Green, 1992) and by calculating the

standard metabolic rate, multiplying it by 2.5, and converting to units of carbon (Schneider

and Hunt, 1982). A Q/B of 36.5 year� 1 is used in the model. Consumption may also be

calculated from the relationship of Nagy (1987): C = 0.495M0.704, where C is daily

consumption in dry grams and M is body mass in grams. A Q/B of 62 year� 1 was used

by Wolff (1994) for northern Chile sea birds, therefore further work probably needs to be

done on the food requirements of New Zealand sea birds. The diet of seabirds is estimated

to be composed of squid, mesopelagic fish, and macrozooplankton, and juvenile fish.

Cherel et al. (1999) show that albatrosses during chick rearing prey on southern blue

whiting juveniles that are 4–5 months old.

Birds have been incidentally killed by fisheries activities on the Southern Plateau.

Bartle (1991) estimated that 1140 white-capped albatrosses were killed during the 1990

southern squid trawl fishery. The main agent of these deaths was the net monitor cable the

use of which was banned in 1992. Grey petrels have also been killed as part of the ling

long-line fishery on the Pukaki Rise. Estimates of bird deaths due to fishing activities have

been made recently for the Auckland Island squid trawl fishery at 38 birds (c.v. = 30%)


(Baird, 1999) and 82 birds were killed in the squid fisheries for 1999–2000 (Buller’s,

Campbell, and white-capped albatrosses, grey and white-chinned petrels, sooty shear-

water, cape pigeon and diving pigeon); about 4% or trawls capture seabirds (Baird, in

press). In the absence of consistently collected data, annual fishing related mortalities of

birds are estimated here to be between 4560 kg WW of white-capped albatrosses or 38 kg

WW of, for example, grey petrels (or 456 or 3.8 kg C). On a per square meter basis over

the whole of the Southern Plateau, these mortalities are 1.05� 10� 6 or 8.76� 10� 9 g C

m� 2 year� 1. In the model, we use the greater value to estimate the potential impact of

fishing on the sea bird populations.

A.2. Seals and sea lions

The New Zealand fur seal (Arctocephalus forsteri) is found on Auckland, Campbell,

Bounty, and Antipodes Islands on the Southern Plateau where they are non-migratory

stocks of a total of about 9600 individuals (Wilson, 1974; Crawley and Warneke,

1979). Adult males weigh 120–185 kg and adult females 40–70 kg (Crawley and

Warneke, 1979). Assuming a 50:50 sex ratio and using a median weight for males and

females, then the average annual wet biomass is 998.4 tonnes. Assuming that these

mammals have the same carbon content as fish (10% of wet weight), the total carbon

biomass is 99.8 tonnes or 0.0002 g C m� 2. This species eats barracouta, cephalopods,

and small fish (Street, 1964). Although southern elephant seals (Mirounga leonina)

have been seen in the New Zealand region, it is assumed that they are not a regular

part of the Southern Plateau ecosystem. Their P/B and Q/B ratios are given by Laws

(1984, Table III).

Hooker’s sea lion (Phocarctos hookeri) is New Zealand’s only endemic seal. Males

weigh about 350 kg and females 110 kg (Gales, 1995), and the total population on the

Auckland and Campbell Islands is probably about 12,500 (Gales and Fletcher, 1999),

although in a recent disease event during January and February 1998, approximately

60% of the sea lion pups and an unknown number of adults died over a 30-day period

on Enderby and Dundas Islands (Baker, 1999). Assuming a 50:50 sex ratio and using a

median weight for males and females, the average annual wet biomass is 2875 tonnes.

Assuming that these mammals have the same carbon content as fish (10% of wet

weight), the total carbon biomass is 288 tonnes or 0.00066 g C m� 2. This species

appears to feed on the bottom and eats cephalopods, crustaceans, and fish and has been

know to eat penguins (Gales and Mattlin, 1997; Childerhouse et al., 2001). The total

average annual seal biomass is therefore estimated to be 0.0009 g C m� 2. Using the

data of (Laws, 1984, Table VIII) and assuming that mortality equals production in a

stable population, then the average P/B for Antarctic seals is about 0.22 year� 1 which

is used here. Annual food consumption in relation to biomass (Q/B) averages 46 year� 1

assuming a daily food intake of 10% of body weight, the upper food intake for captive

animals (Laws 1984); this value is used in the model. Similar values (54 and 47

year� 1) are obtained for fur and Hooker’s seals, respectively, if the equation of Nagy

(1987) for all eutherian mammals is used: C = 0.235M0.822, where C is the daily food

consumed in g DW and M is the wet mass of the animal in g. Jarre-Teichmann et al.

(1998) estimate that Cape fur seals have a Q/B ratio of 19.3 year� 1 and a P/B ratio of


0.95 year� 1 although the former seems too low and the latter too high. The diet of seals

and sea lions is assumed to be composed of squid, and mesopelagic and large fish

(Street, 1964).

In the Bounty/Campbell Plateau region, Hooker’s sea lion are caught incidentally

around the Auckland Islands in the southern squid trawl fishery and New Zealand fur seal

are caught around the Bounty Islands in the southern blue whiting fishery (Baird, 1996,

1997, 1999; Baird et al., 1999; Doonan, 1999). The estimated numbers caught in each

fishery have been averaged over the years that estimates have been made based on

observer data. Therefore an average of 71 Hooker’s sea lions were caught annually from

1988 to 1999 and 119 New Zealand fur seals were caught annually from 1993 to 1998.

This export represents 6.6� 10� 6 g C m� 2.

A.3. Toothed whales and dolphins

There are a number of species of toothed whales and dolphins known to have the

Southern Plateau in their living range (e.g. Gaskin, 1982; Baker, 1990): Arnoux’s

beaked whale (Berardius arnuxii), southern bottlenose whale (Hyperoodon planifrons),

hourglass dolphin (Lagenorhynchus cruciger), Andrew’s beaked whale (Mesoplodon

bowdoini), straptoothed beaked whale (Mesoplodon layardii), spectacled porpoise

(Phocoena dioptrica), goosebeak whale (Ziphius cavirostris), southern rightwhale

dolphin (Lissodelphus peronii), and sperm whale (Physeter macrocephalus). Estimates

have been made of beaked whales (mostly southern bottlenosed whale), but also

including Arnoux’s beaked whale (450,000–800,000 individuals), the hourglass dolphin

(100,000–200,000 individuals) south of 50jS (although these species are distributed

north of this latitude so these are underestimates), and sperm whale south of 30j(128,000–290,000 individuals) (Kasamatsu and Joyce, 1995). Sperm whales are known

to migrate, so their average annual biomass in the Southern Plateau region is estimated

in the following paragraph.

The Southern Plateau extends over 8j of latitude and 16j of longitude. Humpback

whales migrate southwards in spring at an average speed of 15j per month (Brown and

Lockyer, 1984). If we assume that all whales migrate at the same speed through the

region, then the Southern Plateau would be passed in 16 days for a one-way journey

and 32 days for a return journey. Thus, the average annual biomass is calculated based

on 32 days residence. If we assume that whales are evenly distributed with longitude in

the Southern Ocean and that the proportion of the total population that migrates through

the Southern Plateau region is proportional to the longitudinal spread of the plateau

relative to the total extent of the Southern Ocean, then we can calculate an average

annual whale biomass that passes through the Southern Plateau region (16j as a

proportion of 320j, the estimated longitudinal extent of the Southern Ocean) for 32

days by multiplying total whale biomass by 0.0044. Therefore the annual average

biomass of sperm whales is estimated to be 29,604 tonnes wet weight or 2960 tonnes C

or 0.0068 g C m� 2.

The biomass of the other species in the Southern Plateau region are not included. The

surveys of odontocete whales carried out between 1976 and 1988 in spring and summer

in Antarctic waters show that species known to occur occasionally over the plateau have


their main distributions south of 50jS in summer (Kasamatsu and Joyce, 1995). Little is

known of any migration patterns of these species. Therefore, we decided to increase the

average annual biomass of sperm whales by 25%, pending better population estimates of

other species known to occur in the Southern Plateau region. This gives a grand total

biomass of 0.0085 g C m� 2. P/B is calculated from the relationship of Banse and

Mosher (1980): log(P/B) = loga + blogMs, where Ms is in kcal, loga = 1.11, b =� 0.33.

[Note that K. Banse (personal communication) indicated that there is an error in his

Table 2 (Banse and Mosher, 1980): the column labelled a should be labelled loga.]

Whale wet weight is converted approximately to kcal by assuming that like fish, there

are 1000 cal g wet mass� 1 (Schindler et al., 1993). Therefore P/B is 0.042, 0.073,

0.088, and 0.288 year� 1 for an average sperm whale, Arnoux’s beaked whale, southern

bottlenosed whale, and hourglass dolphin, respectively. Therefore, we use a weighted

average of 0.06 year� 1.

Daily food requirements of sperm whales are calculated to be about 3% of body weight

(Brown and Lockyer, 1984) although in other species daily food consumption can vary

from 4% to 14% of body weight. Another method for calculating daily rations, based on

feeding rates of cetaceans in captivity (see Sigurjonsson and Vıkingsson, 1998), is

I = 0.42M0.67, where I is the ingestion rate (kg day� 1) and M is body weight in kg; this

equation gives similar values.

Here we assume that the food requirements dolphins is 4% of body weight. The Q/B

ratio of sperm whales is therefore assumed to be 11.0 and 14.6 year� 1 for hourglass

dolphins. The diet of beaked whales and dolphins is composed of fish (including

demersal fish) and squid (including giant squid) (Brown and Lockyer, 1984; Berzin,

1972).

A.4. Fish (adult)

The total (doorspread) biomass of fish caught on the Campbell Plateau estimated from

Tangaroa and Amatal Explorer surveys from 1989 to 1996 has been very consistent at

about 200,000 wet tonnes (Hurst and Schofield, 1995; O’Driscoll and Bagley, 2001).

Equivalent biomass estimates from two Shinkai Maru surveys during the early 1980s were

about 600,000 tonnes (Hatanaka et al., 1989). The difference appears to reflect differences

in fishing power between the two vessels. The catchability coefficient ( q), which can be

used to scale the estimate up to actual biomass, of most species, is unknown although

modelling suggests that for hoki it is about 0.1–0.15 for the Tangaroa, and 0.25 for the

Shinkai Maru. If a catchability of 0.1 is applied to all species, we get a total standing stock

of all species of about 2.5 million wet tonnes. This 2.5 million tonnes is made up of 1.8%

squid, 9.8% benthic feeders, and 88.4% pelagic feeders (based on proportions in Hurst and

Schofield, 1995).

The main species of fish that have been consistently caught commercially through the

period of data collection on the Bounty/Campbell Plateau are hoki (Macruronus

novaezelandiae), southern blue whiting (Micromesistius australis), hake (Merluccius

australis), ling (Genypterus blacodes), orange roughy (Hoplostethus atlanticus), javelin

fish and rattails (e.g. Hurst and Schofield, 1995), although more than 78 species are

found on the plateau (Anderson et al., 1998). Of these, hake, hoki, ling, and southern


blue whiting are among the quota species for which a Total Allowable Commercial

Catch has been set.

Hoki is the most important fish in the Southern Plateau fishery. The adult fish that

live on the Southern Plateau are part of the Western stock. These fish migrate to the

west coast of South Island to spawn and juvenile fish 2–4 years are found on the

Chatham Rise throughout the year (Annala et al., 2000). There is strong circumstantial

evidence that hoki recruit to the Southern Plateau from the Chatham Rise although the

size and timing of this recruitment is not known. For the purposes of this model, we

assume that there is a fairly steady biomass on the Southern Plateau that is substantially

altered only during the breading season (June to mid-September). In order to estimate

the annual average biomass, we take into account the fact that many adult hoki migrate

out of the area for 3 months (R. Hurst, personal communication) of the year to breed.

Estimations of the total biomass of adult fish (recruited fish z 55 cm) on the Bounty/

Campbell Plateau have been made in November–December 1991, 1992, and 1993

(Chatterton and Hanchet, 1994; Ingerson et al., 1995; Ingerson and Hanchet, 1995). The

mean total doorspread hoki biomass for 1991–1993 on the Bounty/Campbell Plateau

(data from strata 1–5 have been removed) is 77,467 tonnes. 99.8% of this is mature fish

z 55 cm. We assume that hoki biomass is 28.8% of the non-squid biomass of 2.455

million tonnes (see 300–1000 m summer biomass 1990; Hurst and Schofield, 1995).

About 707,040 tonnes of hoki lives on the Bounty/Campbell Plateau in summer.

705,626 tonnes (99.8%) are adult fish and 70% of this migrates to the west coast,

South Island, for 3 months in winter (Livingston et al., 1997). Therefore the average

annual total fish biomass is assumed to be 2,331,516 wet tonnes or 233,152 tonnes C or

0.5377 g C m� 2. In the next section, we calculate the proportion that represents juvenile

fish (0.086 g C m� 2). This is subtracted from the total fish biomass to give the non-

juvenile biomass of 0.4517 g C m� 2.

The annual catch is estimated from data on Quota Management species (Annala et al.,

1999). The average annual catch is about 172,700 wet tonnes (including catches of hoki on

the west coast of South Island where adults went to breed) to which we add an estimated

10% to account for the bycatch species which gives about 189,970 wet tonnes or 18,997

tonnes C or 0.0438 g C m� 2. To estimate total landings, the discards (see below) have

been subtracted to give 0.0430 g C m� 2.

It appears that a large proportion of fish biomass on the Bounty/Campbell Plateau

depends on planktonic food resources. The diet of 7 of the main commercial fish species

was analysed by Clark (1985). He found that hoki, southern blue whiting, and javelin

fish were plankton (water column) feeders, feeding mainly on natant decapods,

euphausiids, amphipods, and some mesopelagic fish. Other dietary information collected

during trawl surveys suggests that arrow squid, hoki, and southern blue whiting are

pelagic or semipelagic feeders with squid feeding on euphausiids, fish, and other squid

and ling feed mainly on other fish and are thought to bottom feeders (Hatanaka et al.,

1989). Using the relative biomass data given by Hurst and Schofield (1995, Table 7),

minus the squid, and the classification of feeding types given above, then 6–11% of fish

biomass (15 major species, rattails, and other quota species) is thought to depend on a

benthic source of food. The diet of hoki on the Campbell Plateau can be divided into

macrozooplankton (16%), mesopelagic fish (23%), fish (52%), and Cephalopoda (9%),


although this can vary considerably with season and location (Clark, 1985, Table 1).

Similarly the diet of southern blue whiting on the Campbell Plateau is 71% macro-

zooplankton and 14% mesopelagic fish, 11% fish, and 4% squid. The diet of javelin fish

is composed of 38% macrozooplankton, 8% mesopelagic fish, 5% fish, 46% Cepha-

lopoda, and 3% macrobenthos. Ling eat 4% macrozooplankton, 12% macrobenthos, and

83% fish, some of which are benthic feeders. Silverside ate mainly macrozooplankton—

salps (84.7%) and macrobenthos (7.4%). The group of fish that have more benthic

organisms in their diets do not feed exclusively on benthic organisms. Smooth rattail ate

17% macrozooplankton, 33% macrobenthos, and 50% fish. Small-scaled notothenid eat

27% macrozooplankton, 33% macrobenthos, and 39% fish. Therefore adult fish are

assumed to eat adult fish, juvenile fish, squid, mesopelagic fish, macrozooplankton, and

macrobenthos.

The annual food consumption/biomass is known only for orange roughy. Orange roughy

adults consume 1.15% of body weight per day and juveniles 0.91% of body weight

(Bulman and Koslow, 1992). Therefore Q/B is 4.2 year� 1 for adults or 3.2 year� 1 for

juveniles. Consumption/biomass is therefore calculated from the empirical relationship

given by Christensen et al. (2000): log(Q/B) = 7.964� 0.204logWl� 1.965T V + 0.083A +

0.532h + 0.398d, where Wl is asymptotic weight of fish (g), T V= 1000/mean habitat

temperature of fish (K), A is the aspect ratio of the caudal fin = h2/s (h is height of tail and s

is surface area of tail), and h and d are a dummy variables expressing food type (0 for

carnivores and detrivores or 1 for herbivores). Southern blue whiting maximum weight

(Wl) of a 60-cm fish is calculated from weight (g) = a� length (cm)b, where a = 0.00515

for females and 0.00407 for males, and b = 3.092 for females and 3.152 for males.

Therefore Wl is 1600 g for females and males. T V= 1000/(7 + 273.15) = 3.57, A= h2/s = 1.42. Therefore Q/B is 2.60 year� 1. The above equation does not apply to eel-like fish

(e.g. hoki, ling). Southern blue whiting is an intermediate-sized fish so its Q/B is used here

for the whole fish compartment.

Annual production/biomass ratios (P/B) for fish is calculated from the equations given

by Banse and Mosher (1980): P/B = aMsb or log(P/B) = loga + blogMs, where loga = 0.44

and b=� 0.26 when Ms is in kcal. The equation for fish is P/B = 2.75M� 0.26. loga = 0.38

when Ms is in g wet weight (Haedrich and Merrett, 1992) or P/B = 2.4M� 0.26. Using this

latter relationship, P/B for 970 g hoki is 0.40 year� 1, for a 90 g hoki is 0.74 year� 1, for

350 g ling is 0.52 year� 1, and for 860 g hake is 0.41 year� 1. Alternatively, by assuming,

in a steady state population, the rate of all mortality (natural and fishing) is equal to

production rate then from population estimates and catch data (Annala et al., 1999), P/B is

estimated at 0.32 year� 1 for southern blue whiting and 0.36 year� 1 for hoki. P/B for the

large fish compartment is set at 0.30 year� 1. Orange roughy P/B ratio has been calculated

to be 0.15 (Pankhurst and Conroy, 1987).

A.5. Discarded fish

Discarded fish are all fish, both target and non-target species that are returned to the sea

as a result of economic, legal, or personal consideration (Anderson et al., 2000). Factory

waste could also be added here, but, because most vessels do not now discard their waste,

this is not included. The calculation of average annual discards is made by using discard


ratios and the total catch of the target species (Anderson et al., 2000; Clark et al., 2000).

Discard ratios have been worked out for arrow squid (0.0171), ling (0.182), southern blue

whiting (0.0159), orange roughy (0.0366), hoki (0.0544), and oreos (0.0286).

A.6. Fish (juvenile)

Juvenile fish are placed in a separate compartment. The proportion of hoki juveniles

< 59 cm total length was on average about 2% by weight across of the total hoki

population in 1991–1993 (Livingston and Schofield, 1996). The proportion of southern

blue whiting juveniles varies widely from dominating the populations as they did in 1993

to forming 14% of the total population in 1995 (Hanchet et al., 1998). These two species

dominate the populations on the Southern Plateau and are fished in a proportion of

approximately 2:1 (southern blue whiting/hoki). Therefore we assume that the biomass of

juvenile fish in the total population is 0.086 g C m� 2. Young fish about 1–2 years old

have a considerably larger P/B (>0.70) than adult fish (using relationships given in

Appendix A.4). P/B of 1 is used. Q/B of 3 is used although a Q/B of 19 has been reported

for North Atlantic cod of 10 cm (Daan, 1973). We assume juvenile fish have a diet of

macrozooplankton and mesozooplankton.

A.7. Baleen whales

The populations assumed to be migrating through the Southern Plateau region are

mainly Eubalaena australis (right whale), Balaenoptera acutorostrata (minke whale),

Balaenoptera musculus (blue whale), Balaenoptera physalus (fin whale), Balaenoptera

borealis (sei whale), and Megaptera novaeangliae (humpback whale). All six species are

thought to breed in tropical, subtropical, or warm temperature waters in winter and feed in

polar or cold temperate waters in summer, with spring and autumn migrations between the

two regions (Brown and Lockyer, 1984). Although it is generally assumed that whales do

not feed on their migration from the Antarctic to the tropics, several species have been

directly observed to feed in the New Zealand region (A.B., unpublished data) if food is

available.

The average annual whale biomass of what passes through the Southern Plateau region

is calculated by the same method used for toothed whales. Therefore the average annual

biomass of baleen whales on the Southern Plateau is 401,170� 0.044 wet tonnes = 17,651

tonnes or 1765 tonnes C (assuming that carbon is 10% of wet weight) or 0.0040 g C m� 2.

Three to four percent of body weight is consumed daily (Brown and Lockyer, 1984 and

references therein) although very young whales have been observed to take up to 13% of

body weight per day. This gives a Q/B of 12.8. We assume that baleen whales eat

macrozooplankton, mesozooplankton, and mesopelagic fish. Production/biomass ratios (P/

B) for some mammals is calculated from the equations in Banse and Mosher (1980) as for

toothed whales. Therefore an 80-tonne blue whale (80� 106 kcal) has P/B of 0.032

year� 1, a 30-tonne humpback whale (30� 106 kcal) has P/B of 0.044 year� 1, a 45-tonne

fin whale (45� 106 kcal) has P/B of 0.038 year� 1, and a 10-tonne minke whale (10� 106

kcal) has P/B of 0.036 year� 1. Sakshaug et al. (1994) also estimate minke whales to have

a of P/B of 0.035. P/B for the whale compartment was set at 0.038 year� 1.


A.8. Squid

The most common squids taken on the Bounty/Campbell Plateau are the warty squid

(Moroteuthis spp.) and arrow squid (Nototodarus sloani). Giant squid (Architeuthis) are

also found in waters 300–600 m deep in this region (Forch, 1998) but their biomass is

unknown. Because giant squid have not been included in squid biomass, the biomass of

this compartment is probably underestimated. A commercial fishery is based on Nototo-

darus sloani. Robertson et al. (1978) record it as part of the mesopelagic fauna. Squid

biomass appears to be about 1.8% of the ‘‘all species’’ biomass (see Hurst and Schofield,

1995, Table 7). Therefore the average annual wet weight biomass of squid is estimated to

be 45,000 wet tonnes. The carbon content of a squid from the Chatham Rise is assumed to

be 20% of wet weight. This is derived from an assumption that ash-free dry weight is

equivalent to the carbon content. Arrow squid dry weight is 22.5% of wet weight and ash

is 6.2% of dry weight (Vlieg, 1988). Vinogradov (1953) gives similar data for dry weight

of Cephalopoda ranging from 13% to 30% of wet weight and ash is 0.9–2.4% of wet

weight. If we assume that ash-free dry weight is equivalent to the carbon content, then the

average annual biomass of squid is 9000 tonnes C or 0.0204 g C m� 2. The annual catch of

squid in the Bounty/Campbell Plateau region was obtained for the years 1992/1993 to

1997/1998 from the Ministry of Fisheries (Trawl Catch, Effort and Processing Return data

and Catch, Effort and Landing Return data). Groomed data on Quota Management species

were compared with the data (Annala et al., 1999) and substitutions made where there was

substantial disagreement. The average annual catch is about 18,000 wet tonnes. Converted

to units of carbon, average annual catch is 0.0042 g C m� 2.

P/B ratios for gonatid squid in the Bering Sea are estimated to be 6.7 (Radchenko,

1992), for Sthenoteuthis pteropus in the tropical Atlantic to be 8.0–8.5 (Laptikhovskij,

1995), and for captive Illex illecebrosus measured to be 2.9–9.1 at 7 jC (Hirtle et al.,

1981). Therefore annual production to biomass ratio for squid on the Bounty/Campbell

Plateau is assumed to be about 8 year� 1 from this work. O’Dor et al. (1980) point out that

growth rates of I. illecebrocsus from field data are well below those for captive animals,

indicating that food supply of the natural population is an important limiting factor. The

daily ration of Loligo pealei ranges from 3.2% to 5.8% of body weight per day

(Vinogradov and Noskov, 1979), which represents a Q/B of 11.7–21.2 year� 1. The mean

daily ration of I. illecebrosus is 5.2% (Hirtle et al., 1981). Here we assume that Q/B is 22

which is necessary if we assume a food conversion rate similar to the highest for I.

illecebrosus (25–36%) (Hirtle et al., 1981). Food of squid appears to be composed of

macrozooplankton (crustaceans), fish, and other squid (Vinogradov and Noskov, 1979).

Here we assume that the diet is made up of adult fish, juvenile fish, squid, mesopelagic

fish, and macrozooplankton (Mattlin and Colman, 1988; Hatanaka et al., 1989).

A.9. Mesopelagic fish

The mesopelagic fauna south of the Subtropical Front is assessed from the work of

Robertson et al. (1978). The most common mesopelagic fish taken in subantarctic water

were the myctophids Protomyctophum normani and Gymnoscopelus procerus. Eight

other mesopelagic species were relatively common: the goniostomatid Cyclothone


pseudopallida, and the myctophids Electrona carlsbergi, Electrona subaspera, Elec-

trona paucirastra, Hygophum hanseni, Symbolophorus ?boops, Lampanyctodes hecto-

ris, and Lampichthys procerus. The sternoptychid Maurolicus australis was a minor

constituent of the subantarctic fauna but was extremely common around shallow areas

of the Chatham Rise. We do not have data on the biomass of mesopelagic fish in the

Southern Plateau region therefore the model calculated this parameter. P/B ratio for

mesopelagic fish is assumed to be about 1. This compares well with data for M.

muelleri for which the growth parameters are very well known (P/B = 1.15) (Ikeda

1996). Similar P/B ratios (0.87–1.38) are given by (Childress et al., 1980) for

mesopelagic fishes off California.

Consumption/biomass ratios have been estimated to be 10.6 year� 1 from the age-

specific daily rations of 0–1.8 year old M. muelleri with a lifetime average of 2.9% or a Q/

B of 10.6 year� 1. Age-specific net growth efficiency of 0–1.8 year old M. muelleri has a

lifetime Q/B average of 16.7 year� 1. Pakhomov et al. (1996) estimated similar Q/B for

subantarctic myctophid species. Here we use 16 year� 1. The diet of M. muelleri is

described by Ikeda et al. (1994) and includes a wide variety of mesozooplankton species,

especially copepods. Therefore we assumed that mesopelagic fish eat mesozooplankton

and macrozooplankton. The biomass of mesopelagic fish is calculated by the model,

assuming that ecotrophic efficiency is 0.95.

A.10. Macrozooplankton

The mesopelagic fauna south of the Subtropical Front is determined from the work of

Robertson et al. (1978). Macrozooplankton are assumed to be mainly Euphausiacea,

although Decapoda and Amphipoda, are also included; Cephalopoda are treated in a

separate group (see above). The biomass of macrozooplankton is unknown in the

Southern Plateau region. Subantarctic open water macrozooplankton biomass ranges

from 0.012 to 4.4 g DW m� 2 and subantarctic around islands ranges from 0.007 to 2.86

g DW m� 2 (Pakhomov et al., 1994). To calculate macroplankton biomass, we assume

that their ecotrophic efficiency is 0.95. Production/biomass ratios are taken from the

literature. Euphausia lucens has P/B = 10.14–16.01 year� 1 (Stuart and Pillar, 1988)

which is high relative to that of Nematoscelis megalops (5–6 year� 1) (Lindley, 1982).

Cartes and Maynou (1998) use P/B ranging from 1.24 to 4.75 for euphausiids and 8.05

for peracarids. Here we use P/B = 10 year� 1 because of lower food availability and

colder temperatures.

Consumption to biomass ratios have been estimated to be 1.205% DW/WW (or about

9% WW/WW) for the mesopelagic shrimp Pasiphaea multidentata (Q/B is therefore

about 33 year� 1) to 0.061% DW/WW for the crab Geryon longipes, with mean values

of 0.364% DW/WW (Q/B 10.7 year� 1) on the middle slope and 0.524% DW/WW (Q/B

15 year� 1) on the lower slope (Cartes and Maynou, 1998). Stuart and Pillar (1990)

show that E. lucens is an omnivore that ingests on a carbon-specific basis 15–60%

phytoplankton, the remainder being mainly small copepods. Five to fourteen percent of

body C day� 1 was ingested by adults and Q/B ranged from 17 to 51 year� 1. Q/B of 33

year� 1 was used. The diet of macrozooplankton (euphausiids) may include phytoplank-

ton, microzooplankton, and mesozooplankton with copepods dominating the diet


(Barange et al., 1991). Therefore we assume that macrozooplankton eat, phytoplankton,

microzooplankton, and mesozooplankton.

A.11. Macrobenthos

Macrofauna biomass on the Bounty/Campbell Plateau is estimated at 0.25 g C m� 2 from

data collected at 750 m on the southern flank of Chatham Rise (S.D.N. et al., unpublished

data). A P/B ratio for macrofauna can be estimated from the relationship given by Brey and

Gerdes (1998) showing an exponential increase of annual community P/B with water

temperature. If a mean bottom temperature of 6.5 jC for 600 m is assumed (Livingston and

Schofield, 1993; Schofield and Livingston, 1994), and the regression equation of Brey and

Gerdes is applied, an annual P/B of 1.00 is obtained. A P/B ratio of 1.83 is used by Cartes

and Maynou (1998) for polychaetes. We assume that production/consumption is 0.35.

The macrobenthos are mainly deposit feeders, for example, on Chatham Rise, and

average of 65–71% of polychaetes were deposit feeder (Probert et al., 1996; P.K.P.,

unpublished data). Since polychaetes dominate the macrofaunal, their trophic structure

should be a reasonable representation for the macrofauna as a whole. We have assumed

that the macrobenthos is fuelled largely by the sediment bacteria but also feed on other

benthos although benthic crustaceans have been shown to also eat euphausiids (Cartes and

Maynou, 1998). In subantarctic water, 7–22 Ag Chl m� 2 day� 1 reaches 550 m (Nodder

and Gall, 1998) but this is a very small quantity and is ignored as a source of food for the

macrobenthos. Therefore we assume that the macrobenthos eats macrozooplankton,

macrobenthos, meiobenthos, and bacteria.

A.12. Mesozooplankton

The average annual biomass (wet weight and as carbon) of mesozooplankton is

calculated using data collected in 1993 (Bradford-Grieve et al., 1998), and historical data

collated by Bradford (1980). These data have been adjusted for the average depth of the

water column over the Plateau. Mean annual carbon biomass 0–615 m from these data is

1.058 g C m� 2. The production/biomass ratio for mesozooplankton for low productivity

water is about 12 (Shushkina et al., 1998). This may be compared with P/B of a subtropical

copepod Acrocalanus inermis which was measured by Kimmerer (1983) and varied from

0.07 to 0.36 day� 1 and 0.2 day� 1 (Vidal, 1980). Baird and Ulanowicz (1989) estimated an

average P/B ratio of 0.37 day� 1 over an entire year in Chesapeake Bay, an enclosed coastal

system. Secondary production is not continuous in subantarctic water because primary

production is very low in winter (Bradford-Grieve et al., 1997). Therefore P/B is estimated

by assuming that secondary production occurs over only 6 months of the year and daily P/B

is 0.11. We assume that P/B for mesozooplankton is about 20 year� 1.

Food intake has been determined experimentally (see Parsons et al., 1984) and ranges

from 10% to 20% of body weight per day for large crustaceans to 40–60% per day for small

crustaceans. Paracalanusmay eat 1.5 Ag N Ag body N� 1 day� 1 (Checkley, 1980) although

their specific ingestion of C was 3.6 day� 1 when feeding on N-deficient Thalassiosira. For

large copepods such as C. finmarchicus, Ohman and Runge (1994) showed that, in the

lower estuary region of the Gulf of St Lawrence, total food was ingested (diatoms


dominant) at the rate of 42–48% of body C day� 1 and in the open gulf total food was

ingested (dominated by aloricate ciliates) at a rate of up to 4% of body C day� 1. At all these

stations the copepods were laying eggs although the authors consider the possibility that

these copepods might not have been in equilibrium with the food supply. The implication

appears to be that protozoa may be a much better food source that autotrophic food

particles. It was assumed that P/Q is 0.35. We assume that the mesozooplankton feed on

phytoplankton, microzooplankton, and mesozooplankton (Bradford-Grieve et al., 1998;

Zeldis et al., 2002). Assimilation of copepods is assumed to be 0.3 for animals that are

feeding on microzooplankton (Pavlovskaya and Zesenko, 1985).

A.13. Ciliates

The average annual biomass of ciliates (as carbon) is calculated using data collected in

a number of months (Bradford-Grieve et al., 1998; Hall et al., 1999; J.H., unpublished

data). Integrations are made to 100 m and the assumption is made that there are no ciliates

below 100 m if there are no measurements below this depth. Ciliate carbon biomass was

calculated using a factor 0.19 pg C Am� 3 (Putt and Stoecker, 1989). The average annual

ciliate biomass is 0.167 g C m� 2.

Mean daily P/B of ciliates is 0.24 (n = 5) calculated from dilution grazing experiments

(J. H., unpublished data). These data are from subantarctic waters in August and January–

February; there was little difference in P/B between the two periods. A ciliate production

rate of 0.3 day� 1 (110 year� 1) is near the mean of estimates from a number of studies

tabulated by Kiørboe (1998) although growth rates of up to 0.9 day� 1 have been measured

(Verity et al., 1993). Also, in the subarctic Pacific, ciliate production of 0.10 day� 1 is

given by Landry et al. (1993) although this may be too low if predators were not fully

excluded from incubations. We therefore use an annual P/B of 88. We assume that

production/consumption is 0.36. The proportions in which ciliates consume their food

(phytoplankton and heterotrophic flagellates) can only be estimated although we know that

ciliates consume 70% of the biomass of heterotrophic flagellates and autotrophic biomass

per day (J.H., unpublished data).

A.14. Heterotrophic flagellates

The average annual biomass of heterotrophic flagellates (as carbon) is calculated using

data collected in a number of months (Bradford-Grieve et al., 1998; Hall et al., 1999; J.H.,

unpublished data). Integrations are made to 100 m and the assumption is made that there are

no heterotrophic flagellates below 100 m if there are no measurements below this depth.

Heterotrophic flagellate carbon biomass was calculated using calculated cell volumes

(Chang and Gall, 1998). The average annual heterotrophic flagellate biomass is 0.307 g C

m� 2.

Mean daily P/B of heterotrophic flagellates is 0.80 (n = 10) (292 year� 1) calculated

from dilution grazing experiments (J.H., personal communication). These data are from

subantarctic waters in August and January–February; there was little difference in P/B

between the two periods. Growth rates of heterotrophic microflagellates of >2 day� 1 have

been measured when conditions are not limited by iron (Chase and Price, 1997) but are


< 1 day� 1 at the low prey Fe/C of 9 Amol mol� 1 observed in the open subarctic Pacific

(see Tortell et al., 1966). In low iron growth conditions, carbon-specific growth of

microflagellates was 0.7–1.6 day� 1. The lower end of these growth rates is similar to

the growth rates calculated for subantarctic waters from dilution grazing experiments. We

assume that production/consumption is 0.35.

The proportions in which heterotrophic flagellates consume their food (bacteria and

phytoplankton) can only be estimated. We know that heterotrophic flagellates consume

4.4% of picophytoplankton biomass and 2.4% of bacterial biomass per day (Safi and Hall,

1999; J.H. unpublished data). Assimilation efficiency ((ingestion� excretion)/ingestion)

of heterotrophic flagellates in low iron conditions is 0.84 (Chase and Price, 1997) although

we initially use 0.70.

A.15. Meiobenthos

Meiofaunal biomass for the Bounty/Campbell Plateau at slope depths is estimated from

some nearby measurements (S.D.N. et al., unpublished data) and values from the literature.

Meiofaunal biomass integrated to 5 cm at 750–1000 m depth just north of the plateau

ranges from 0.07 to 0.09 g C m� 2. These data are somewhat lower than those derived from

regression equations for the temperate North Atlantic which indicate values of 0.35–0.40 g

C m� 2 for a depth of 600 m (Tietjen, 1992; Soltwedel, 2000). On the basis of various

regressions of density vs. depth (mainly North Atlantic), the total meiofaunal density at 600

m may be calculated to be f 1400 individuals 10 cm� 2. A mean weight per individual

nematode (the dominant taxon) of 0.2 Ag dry weight, or 280 Ag dry weight 10 cm� 2,

converts the number of individuals to 1.12 g wet weight m� 2, or 0.112 g C m� 2, assuming

dry weight to be 25% of wet weight and carbon to be 40% of dry weight (Feller and

Warwick, 1988). These data are of a similar order of magnitude therefore we assume an

average annual meiofaunal biomass of 0.1 g C m� 2. Annual P/B ratios of meiofauna vary a

lot, but 10 is often taken as an average value (Feller and Warwick, 1988). The prime source

of food for the meiobenthos is assumed to be bacteria with some contribution from other

meiobenthos. Annual production/consumption was assumed to be 0.31.

A.16. Bacteria (water column)

The average annual biomass of bacteria (as carbon) is calculated using data collected

just north of the region (Bradford-Grieve et al., 1998; Smith and Hall, 1997; J.H.,

unpublished data) and is estimated at 0.598 g C m� 2 using the carbon conversion factor of

Fukuda et al. (1998). This value is likely to be an underestimate as we have not taken into

account the likelihood that bacterial biomass is greater around the islands due to the

additional input of detritus from macroalgae as well as from the water column system

because of greater phytoplankton biomass and consequent trophic flows. The annual

production/biomass ratio for bacteria is estimated to be 87.4 year� 1. Shushkina et al.

(1998) estimate bacterial P/B to be 92 based on the analysis of for low productivity waters

whereas Sorokin (1999, Table 2.2) gives P/B of 0.5 day� 1 for eutrophic coastal habitats;

0.6 day� 1 in mesotrophic temperate seas; and 1.2 day� 1 in oligotrophic tropical seas

which seem extremely high. Estimates of production/consumption (P/Q) where obtained


by Tortell et al. (1966) for iron-limited water in the North Pacific. These estimates were

0.12–0.18, although we have used 0.23.

A.17. Bacteria (sediment)

We do not have direct measurements of bacterial biomass for the Bounty/Campbell

Plateau. Data from the surface 3 cm are available at slope depths on the southern flank of

Chatham Rise (S.D.N. et al., unpublished data). In order to compare these data with those

from other studies where sediment bacteria are integrated to greater depths, we use the data

at one station in summer which indicate that biomass at 0–9 cm is 3.6–4.5 times the

biomass at 0–3 cm and bacterial production at 0–9 cm was 1.2–1.5 times the production

at 0–3 cm. These data are very variable and because we suspect that the Chatham Rise

sediment is strongly influenced by the Subtropical Front sitting above it, we have decided

to use a biomass estimate from the regression of Deming and Yager (1992) of about 1.5 g

C m� 2 (to a sediment depth of 15 cm).

Annual bacterial production (0–3 cm) ranges from 0.18 to 0.35 g C m� 2 year� 1 for

stations 750–1000 m and these same stations have an annual bacterial P/B ratio of about 1

(S.D.N. et al., unpublished data). These values are considerably lower than the average

bacterial production of 16.9 g C m� 2 year� 1 reported by Kemp (1994) for slope sediments

( < 2000 m). Alongi (1990) found bacterial productivity (surface 0–5 mm sediment) to

decrease significantly with depth (695–4350 m) and, from the regression given, one gets a

figure for 600 m depth of 34.7 g C m� 2 year� 1. For the Celtic Sea (135–1680 m), Poremba

and Hoppe (1995) measured microbial activity in the upper 10 cm of sediments. Activity

was always very much higher in the top 0–1 cm of sediment. They give specific growth

rates for their station at 571 m: bacterial P/B in the top 0–1 cm was 0.0298 day� 1 or 10.9

year� 1. Alongi (1990) also gives specific growth rates for bathyal and abyssal stations.

They vary widely from 0.001 to 0.12 day� 1 (0.37–43.8 year� 1) but decrease with water

depth. Sorokin (1999) gives annual P/Bs of 14.6 and 7.3 off Japan. Using these latter values

of biomass and production, one gets average annual P/B ratios of around 11–12. Because of

this uncertainty, P/B is calculated by the model by assuming that ecotrophic efficiency is

0.80. A growth efficiency (P/Q) of 0.3 (Kirchman, 2000) is assumed here.

A.18. Phytoplankton

Average annual biomass is estimated to be 0.895 g C m� 2 from Bradford-Grieve et al.

(1999). This value does not take into account the often heightened chlorophyll concen-

trations around the islands and over the Pukaki Rise (Murphy et al., 2001). We therefore

increase annual average phytoplankton biomass by a factor of 1.2 to give 1.07 g C m� 2.

Daily primary production of 28–62 mg C m� 2 day� 1 in winter (June) and of 230–271

mg C m� 2 day� 1 in early spring (October) in SAW was measured in 1993 (Bradford-

Grieve et al., 1997), although in October, 1997, primary production in SAW (390–680 mg

C m� 2 day� 1) was greater than in 1993 (Boyd et al., 1999). As it is difficult to be sure of

the annual average primary production in the region from these data, we investigated

estimates from satellite data. Annual primary production, estimated from the data

presented by Moore and Abbott (2000) for the Subantarctic Water Ring, is about 80 g


C m� 2 year� 1. Therefore, these data give a P/B of about 89 and 118 year� 1, respectively.

P/B of 89 year� 1 is very low compared with annual P/B = 218 given by Shushkina et al.

(1998) for low productivity water.

It is noted by Ducklow (2000) that: ‘‘bacterial production in the water column is

maintained in a remarkably constant ratio to primary production, averaging about 0.15–

0.20 across oligotrophic and oceanic HNLC and upwelling and blooming systems.’’ For

The Southern Plateau system, however, the ratio is considerably higher at 0.66 (taking

bacterial and primary production as 53 and 80 g C m� 2 year� 1, respectively). Our

bacterial biomass and production estimates are typical. This suggests that annual primary

production over the Southern Plateau might be nearer to 265 g C m� 2 year� 1 and annual

P/B might be about 248. Nevertheless, Southern Plateau phytoplankton production thus

estimated is much lower than the range of values given by Ducklow in his Table 5 (465–

1548 mg C m� 2 day� 1).

A.19. Detritus and dissolved organic carbon (water column)

Annual average integrated particulate carbon (PC) for the water column is estimated

from data collected in 1993 (spring and winter) (McCarter and Hall, 1996a,b). Where

only particulate nitrogen (PN) data were available, particulate carbon was estimated

assuming that PC/PN is 6. Data were integrated down to 615 m. Detritus was estimated

by subtracting phytoplankton, bacteria, and microzooplankton biomass from the PC

data.

Antarctic concentrations of dissolved organic carbon (DOC) in spring surface water of

the Atlantic sector of about 0.46–0.66 mg C l� 1 or g C m� 3 (Kaehler et al., 1997) and

0.52–0.84 g C m� 3 in October in subantarctic water east of New Zealand (M. van Kooten,

personal communication). In February–March in the Atlantic sector of the Antarctic,

Romankevich and Ljutsarev (1990) record surface concentrations of DOC >1.2 mg C l� 1.

Therefore we assume that annual average concentrations of DOC on the Southern Plateau

of 0.9 g C m� 3 and that these concentrations occur to 615 m to give an integrated

concentration of 554 g C m� 2. The sum of the particulate detrital carbon and DOC is

17.6 + 554 = 572 g C m� 2.

A.20. Detritus and dissolved organic carbon (sediment)

Average annual organic carbon content of the subantarctic sediment on the Pukaki Rise

is estimated from Carter et al. (1999). At 28 cm below surface, the total organic carbon

content of the sediment is 0.28% of the dry weight of sediment. The dry density of sediment

at 3.10 m below the surface is 1.031 g cm� 3. Therefore assuming the properties of the

sediment about the sampling depth are similar, 1 m2 of sea floor down to 15 cm depth is

150,000 cm3 or 154,650 g. Therefore, the carbon content of 0.28% of dry sediment is 433 g

m� 2. This result may be compared with data from 450 m on the south side of Chatham

Rise. At this station, the surface 5 cm of sediment has 0.6% total organic carbon. The dry

density of the sediment at this depth is about 1.65 g cm� 3. One square meter of sea floor

down to 15 cm depth is 150,000 cm3 or 247,500 g. Therefore, the carbon content of 0.28%

of dry sediment is 693 g m� 2. The actual average figure for the Plateau is probably


somewhere between although we have chosen the lower value since the water column input

to the sediment is known to be low (H. Neil, personal communication). Annual average

detritus supplied to the sediment has been measured by Neil on the southern flank of Pukaki

Rise at 50j15.64VS, 171j28.35VE to be 1.23 g C m� 2 year� 1.

We assume pore-water has similar concentrations of DOC as are found at Station

M271-96 at 806 m SW of Britain at about 48jN in summer (Otto and Balzer, 1998) and

the porosity of the sediment ranged from 0.66 to 0.47 (W. Balzer, personal communica-

tion). The total DOC down to 15 cm is 0.43 g C m� 2; carbon in this form is assumed to be

included in the Total Organic Carbon content of the sediment of 433 g C m� 2.

A.21. Assimilation efficiency

The portion of food consumed and not assimilated was taken as 0.2 (the default for

Ecopath) except for mesopelagic fish, macrozooplankton, macrobenthos, mesozooplank-

ton, ciliates, heterotrophic flagellates, meiobenthos (set at 0.4, 0.4, 0.4, 0.35, 0.3, 0.3, 0.3),

respectively, mainly to ensure there is enough substrate for bacteria in the model.

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