Oxygen consumption in Palaemon pacificus (stimpson) (Decapoda: Palaemonidae) in relation to temperature, size and season

Camp. Biuchcm.

Phwiol.

030&9629/X5 $3.00 + 0.00 ‘(‘81985 Pergamon Press Ltd

Vol. XIA, No. I, pp. 71&7X.1985

Printed rn Great Britain

OXYGEN CONSUMPTION IN PALAEMON PACIHCUS (STIMPSON) (DECAPODA : PALAEMONIDAE) IN RELATION TO TEMPERATURE, SIZE AND SEASON W. D. EMMERSON* Department

of Zoology,

University

of Port Elizabeth.

P.O. Box 1600, Port Elizabeth.

South Africa

(Received 27 July 1984) Abstract-

1. The effect of temperature, body mass and season on the oxygen consumption of the shrimp Palaemon pac~jicu.s was investigated. 2. Temperature-specific oxygen consumption values between 10 and 30°C varied from 0.078 to 0.735 mg 0, g ‘hr ‘, similar to published values for shrimp. 3. There was no significant difference between summer and winter temperature-specific regressions indicating no seasonal acclimation. 4. The predictive respiratory equation combining shrimp size (w. temperature (T) and season is R=0.00255 T-“” ,W’““or in terms of energy (E. Jhr ‘10 ‘), E=0.03606 T~)7’W”‘1. 5. A marked increase in oxygen consumption was measured during ecdysis. 6. Q,,,values varied with temperature but not with mass or season. ranging from 2.981 at 30°C to 4.695 at 15°C.

INTRODUCTION

each container monitored for 15 min/hr. The containers used were glass tubes. The water was maintained at the described temperature ( f O.l”C) using thermoregulators and room temperature control. Flow rates through each container ranged from 18 to 23 ml/min. were checked at least twice per day and varied less than 5% between individual readings. The water bath holding the experimental chambers was covered with an opaque plastic sheet to prevent the shrimp being disturbed yet still allowing light penetration. The photoperiod was 12 : 12 (L D) in all cases. Monitoring commenced after the shrimp were placed into the respirometer, but readings were only used after a 5 hr settling period. Depending on size class availability P. pac~$cus were either collected from tidal pools at Pollock Rock. Port Elizabeth (34”OO’S. 25”3O’E) or from Zostera capensis beds in the Swartkops estuary, Port Elizabeth during winter (JulySeptember, 1980) and summer (December-February, 1981). Fresh shrimp were used for each experiment after an acclimation period of 24 hr to the test temperature during which time they were not fed. To determine the effects of temperature. size and season on the metabolic rate, the oxygen consumption of a size range of shrimp was measured over at least 36 hr at 10. 15. 20 and 25°C in winter and 15, 20. 25 and 30°C in summer. This temperature range (1&3(K) was chosen as it corresponds to the temperature to which P. paczficus is naturally exposed in its wide distribution. This shrimp occurs from cold west coast waters (Cook and Achituv, in press) through the temperate eastern Cape coast to sub-tropical systems in Natal (Millard and Harrison, 1954; Robertson, 1984). Fresh filtered seawater at 35”/,,S was used for all experiments, After each experiment the shrimps were removed. oven dried at 60°C for 24 hr and weighed to the nearest mg. Oxygen consumption was determined as follows:

consumption

has been widely studied in Crustacea and has been shown to be affected by environmental factors such as temperature, salinity, season, light and oxygen concentration (Lofts, 1956; McFarland and Pickens, 1965; Kinne. 1970; Bridges and Brand, 1980) and intrinsic factors such as body size, activity, moult stage, starvation, sex. haemocyanin level and biological rhythms (Wolvekamp and Waterman, 1960; Subrahmanyam, 1976; Silva and Regnault, 1980; Alcaraz and Sarda, 1981; Hagerman and Weber. 1981; Du Preez, 1983). In southern Africa most metabolism studies have been confined to sandy beach macrofauna (Hanekom, 1975; Dye, 1979, 1980; Brown and da Silva, 1979; Dye and McGwynne, 1980; Du Preez, 1983). with a smaller contribution on estuarine macrofauna (Marais, 1980: Hanekom. 1980; Dye and van der Veen, 1980; Els, 1983). This paper investigates the effect of temperature, size and season on the oxygen consumption of the palaemonid shrimp, Paluemon pact$icus which is ubiquitous in estuarine systems, tidal pools and the nearshore environment of the Cape, South Africa (Emmerson, 1983). Routine oxygen consumption. defined as the mean oxygen consumption rate over a period of 24 hr, is used here as this is the most useful measure of bioenergetic studies (Hart, 1980). Oxygen

MATERIAL AND METHODS Oxygen consumption was measured in a continuous respirometer as described by Marais et crl. (1976) Emmerson and Strydom (1984) using a Radiometer base analyser and chart recorder. The system has channels (three experimental and one control chamber)

*Present address: Department of Zoology, University Transkei, Private Bag X 5092, Umtata. Republic Transkei. Southern Africa.

flow and acidfour with

R = APO, (chart divisions) x Aow rate x O2 value of I chart division

(1)

where R = pg 0,consumed hr ‘; APO, = the number of chart divisions deflected from control; flow rate=mlhr ’ and 0, value of 1 chart division = 0, solubility in mgl~ ’ at T(“C)/ number of chart divisions between zero and preset saturation level for given temperature, pressure and salinity. The chart was usually set on 100 divisions for saturation at any given temperature and pressure and then brought down by

of of

71

72

W.

salinity factor. e.g. 0.832 for 35”/,,S. ration = 83.2 divisions. the

D.

EMMERSON

i.e. satu-

RESULTS

Oxygen consumption varied during the experimental periods, although no repeatable pattern was evident (Fig. 1). The mean oxygen consumption values for each experimental run were therefore used and found to be curvilinearly related to dry mass at each experimental temperature in the form R=a,

W’.

(2)

where a, and b, are constants, R is the oxygen consumption (ugO> hr-‘) and M is the body drymass (mg). Equation (2) may be log,,, transformed to yield the double logarithmic expression log,,, R = log,,, a, + b, log,, M. Equation (2) may also be expressed requirements (Hart. 1980)

(3)

in terms of energy

E = a2 W’.

(4)

where E is the energy requirement of metabolism in joules (J), a? = Q,,a, where QO.is the energy equivalent of oxygen consumption (Jmg 0,-r) and b,= b, the

slope of the linear regression. As the Q,, for P. pac$cus is not known, the general herbivore value of 14.14 JmgO,~‘(Elliot and Davidson, 1975) was used. The relationship between log R and log Mat 10, 15, 20 and 25°C (winter) is shown in Fig. 2(a) and at 15. 30, 25 and 30°C (summer) in Fig. 2(b). The temperature specific regressions for winter and summer acclimated P. pacificus are given in Table 1. Analysis of covariance (Zar. 1974) showed that the slopes for each temperature were not significantly different during summer or winter. but the elevations for each temperature were highly significantly different (Table 1). Further paired comparisons (TV-test, Zar. 1974) between different temperatures during winter and summer also showed highly significant differences in temperature-specific elevations (Table 2). No signiticant difference either in slope or in elevation between the same temperatures during winter and summer (Table 2) showed that season had no effect on metabolism. A single expression relating oxygen consumption R or energy of metabolism. E to size. M, temperature. T(C) and season is necessary in energetic studies where the relationship of a, to temperature and season is required using a common slope. b,. There was no significant difference in intercepts with season 1

c-7

0 x

1:;

25%,

llOm9

11

I

3

20°c,

145

mg

-II 20%.

‘1-i

70mg

1 I

L

II,““‘f

,,,,,,,,,,,,,,,,,,,,,,,I,II,,,,,II

18hOO

Fig.

24hOO

06hOO

i2hOO

I. Example of individual P. pacificus oxygen consumption low tide times arc indicated. An example of a moultincg

24hOO

16hOO with shrimp

time. is also

Bar

12hOO

06hOO represents

given.

night.

E denotes

High

ccdysis.

and

Oxygen

13

I

I

I

1.0

0.5

in Palaemon

consumption

I

I

2.0

1.5

2.5

3.0

SUMMER q 30

1.5

LOG ,. M

I

I

I

I

1.0

2.0

Img

dry

? 0

2.5

mass1

Fig. 2. Oxygen consumption in P. pucifcus during A winter (4 temperatures) and B summer (4 temperatures) in relation to shrimp size. Equations for the temperature-specific lines are given in Table 1.

(Table 2) but a curvilinear increase of a, with temperature was found (Fig. 3) described by a I ~0.00’55

T237’4’

(5)

(r=O.993; N=8; P
ficantly different (Table 1). A predictive equation is then obtained if the common slope b, is used and equation (5) is substituted for a, in equation (2) R=0.00255 Calculated

summer

70.‘72~832.

(7)

and winter temperature-speci-

Table I. Analysis ofcovariance on winter and summer acclimated Paluemon po~+/icus to indicate temperature-specific differences in slopes(b) and elevations (a) in the relationships between oxygen consumption (R. mgOz hr ‘) and dry mass (M, mg). NS, not significant: S”, highly significant Elevation

Slope (b) Season

Winter

Summer

.Y

(“C)

I 2 3 4

IO IS 20 25

5 6 7 8

IS 20 2s 30

log,,R=log,,a,+b,

log,&

N

I

-0.1752+0.8544log,,M 0. I393 + 0.8674 log,&4 0.5183 +0.7890 log,,@ 0.7710+0.7801 log&f

I? I6 24 IO

0.933 0.981 0.944 0.962

0.1409+0.8764log,,M 0.5207+0.8192 log&I 0.7050 + 0.8229 log,,M 0.8873 f0.8463 log,@

IO IO II IO

0.992 0.976 0.9x3 0.9X6

(a)

Comparisons

F

r

P

F

r

P

l/2/3/4

I .6X3

3: 54

NS 0.776

63.699

3 : 57

P/c** c[ 0.00001

5/6/7;8

0.276

3 : 33

NS 0.848

91.542

3 : 36

s** ~0.00001

74

W. D. EMMEKSON

Table 2. Comparisons within seasons (summer and winter) and between seasons (same temperatures) to test differences in the temperature-specific relationships between oxygen consumption (R, ug 0, hr ‘) and dry mass (M. mg) in Puluemon pacz&u~ Slope (b) Comparisons

Winterdifferent

N

between temperatures

i

I

I’

NS

- 5.072

25

2,:3

NS

-5.181

37

-4.138

31

~~

NS

~

NS

-5.996

17

L 617

NS

-4.559

18

7/g

NS

-6.027

1X

-0.701

23

- 1.039

31

-0.699

18

2/s

.

t

516

Season-between same temperatures

8-

P

I!‘2

3,‘4

Summerbetween different temperatures

\

Elevation

0.003

NS 0.501 NS 0.415 NS 0.495

32

3,‘6

-0.216

30

417

-0.013

17

(a) P s** 0.00002 s** < 0.00001 s** 0.00012 s** 0.00001 s** 0.00012 s** 0.0000 I NS 0.245 NS 0. I53 NS 0.246

Sunlmer

0 winter 6-

4-

2-

/CI

1;

14 I

,‘rj

1’8

2’0

212

24I

26I

2’8

30I

T’C Fig, 3. Intercepts

(a) from summer

and winter temperature-specific

regressions

(Table I) as a function of

temperature. fit values of metabolic energy requirements for a representative size spread (10, 50 and 300mg drymass) of shrimp was compared with estimates from the predictive equation (7) (Table 3). Deviation was found to be generally less than 10% with the greatest deviations occurring at the lowest temperature and the largest mass, e.g. - 20.8% for a 300 mg shrimp at 10°C. The effect of temperature. mass and season on Q,, (calculated from the temperature-specific regressions and not the predictive equation 6) is shown in Table 4. While QIOappeared to be unaffected by mass. there was a tendency for Q10to increase with a decrease in temperature during summer and winter. Values ranged from 2.981 to 4.526. with an overall mean of 3.704 (Table 4). Shrimp which moulted during an experiment showed a marked increase in oxygen consumption. An example is given in Fig. I. Oxygen consumption increased sharply 4-5 hr before ecdysis to reach a maximum just prior to ecdysis when it was approxi-

mately double the previous level. The oxygen consumption subsequently declined over the following 12 hr. The moult occurred at night. A number of crustacean respiration studies have been expressed as specific oxygen consumption rates using fresh mass (McFarland and Pickens. 1965: Kutty et al.. 1971). In order to compare data directly with those of other authors, it was necessary to calculate temperature-specific oxygen values (ugOZ hr-‘) in terms of specific oxygen consumption (mg0, g-’ hr-I). Dry mass values for P. pacz$?us were converted to fresh mass values from the linear relationship M,,= 1.738 + 0.265 M,,

(8)

(N = 80; r = 0.996; P < 0.01; unpublished data) where Md is dry mass (mg) and M, is fresh mass (mg). Specific oxygen consumption values for a representative size range (31, 182 and 263 mg fresh mass) of shrimp are given in Table 5.

Oxygen

consumption

Table 3. Estimates of the metabolic pacificus of given dry mass at various the temperature-specific Temperature T (“C)

Season W W W W W W S s s W W W S S S W W W S S S S S S

10 10 IO 15 15 15 15 15 15 20 20 20 20 20 20 25 25 25 25 25 25 30 30 30

Table

4. Summary

of Qlo values

energy requirements (J.hr’ x 10 ‘) for Palaemon summer and winter temperatures as determined from regressions and the predictive equation

Dry mass, M (mg) ___-

Temp. specific regression

10

61.5 261.2 1235.2 143.6 580.1 2744.5 147.1 603.0 2899.4 286.9 1021.4 4199.1 309.3 1155.9 50 16.0 513.6 1802.3 7292.0 476.8 1792.9 7832.1 765.6 2988.9 13615.3

50 300 10 50 300 IO 50 300 10 50 300 10 50 300 IO 50 300 10 50 300 10 50 300

of

75

in Palaemon

P. pucifitu~ at different

temperatures dry mass (mg)

Mass

Predictive equation

% deviation

51.7 220.3 978.2 151.1 576.4 2559.6 151.1 576.4 2559.6 299.0 1140.7 5065.0 299.0 1140.7 5065.0 507.6 1936.7 8599.9 507.6 1936.7 8599.9 782.3 2984.8 13253.9

for winter

- 14.5 - 17.5 - 20.8 + 5.0 - 0.6 - 6.7 + 2.6 - 4.4 -11.7 + 4.0 + 10.4 + 17.1 - 3.3 - 1.3 + 1.0 - 1.2 + 6.9 + 15.2 + 6.1 + 7.4 + 8.9 + 2.1 - 0.1 - 2.6

and summer.

Shrimp

(mg)

T(‘C)

IO

50

300

Meall

WI5

4.526

4.342

Sl5

4.359

4.513

4.444 4.695

4.437 4.522

\ 1

4.480

w20

3.996

3.521

S20

4.205

3.834

3.060 3.460

3.526 3.833

\ 1

3.680

W25

3.580

3.529

3.473 3.123

3.517 3.103

\ I

3.315

3.734 3.477

3.344 3.341

3.683

3.704

S25

3.083

3.102

w30

2.981

3.317

s30

3.211

3.334

Meall

3.743

3.6X7

DISCUSSION

Increase in oxygen consumption or decrease in specific oxygen consumption with size is well known in Crustacea (Zeuthen, 1953; Bertalanffy, 1957: Subrahmanyam, 1962; van Donke and de Wilde, 1981). Slopes (b) range from approx. 0.67 to 1.0 (Zeuthen, 1953: Wolvekamp and Waterman, 1960) but are generally less than 1.0 (Bridges and Brand, 1980) averaging around 0.85 for Crustacea (Weymouth et a/., 1944). Slopes for P. pac$cus varied from 0.780 to 0.876 with a mean b value of 0.832, similar to the average value for Crustacea. The absence of an obvious increase in oxygen consumption with a 24 hr periodicity indicates that there is no circadian rhythm, while no repeatable peak at 12 hr further suggests there is also no tidal rhythm. A tidal rhythm may be expected as P. pacifi-

..~

3.343

cus exhibits a tidal migration pattern in Zostera capensis beds, with an associated feeding rhythm (unpublished data), while recruitment of this shrimp into tidal pools is also dependent on tide. Both Palaemon elegans and Palaemon serratus from tidal pools show tidal activity rhythms which occur on the “expected” ebb and lightldark changes (Rodriguez and Naylor, 1972). Tidal migration may be passive in P. pacz~cus (see Emmerson, 1983) so an increase in oxygen consumption is not illicited. However. even surf zone occupants like Ovalipes punctatus (Du Preez. 1983) and Gastrosaccus psammodytes (Dye. 1980) do not display tidal (12 hr) rhythms in oxygen consumption but circadian (24 hr) ones. Hart (1980) also found no die1 pattern was exhibited by Caridina nilotica. Subrahmanyam (1976) did, however, find tidal and diurnal rhythms in activity and oxygen consumption in Penaeus duorarum but this is to be

-I6

W. D. EMMERSON Table 5. Oxygen consumption (fig O2 hr ‘) and specific oxygen consumption (mg 0: g ’ hr ‘) of faiaemun pat@-us of given freshmass at various summer and winter temperatures as determined from the temperature-specific regressions

.~.

mgO,g

Season

U”C) ‘hr ’ IO III IO 15 15 I5 15 15 IS 20 20 20 20 20 20 25 25 25 25 25 25 30 30 30

Fresh mass (mg)

W W W W W w s s s W W W s s s W W W s s s F s S

Mean 20

4.774 18.897 87.355 IO.156 41 .OZh lY4.045 10.403 42.645 x5.050 20.290 77.235 ‘96.966 21.874 Xl.747 354.73x 36.232 127.461 515.700 33.720 126.796 553.897 54.144 211.379 ‘)67.X93

0.153 0.104 0.07x 0 326 0.225 11.I?2 0.332 0.234 n.lHZ I).650 0.397 0.264 0.70 I 0.449 0.3 is 1,164 U.70# 0.45x I.081 0.696 0.492 I.735 I.161 0.X56

446.3

170.190

0.539

expected as this species buries itself during the day, emerging at night to feed. Specific oxygen consumption values for P. pacificus between 10 and 30°C varied from 0.078 to 1.735 mg0, g-’ hr-‘. respectively (Table 5). Oxygen consumption per gram body mass was higher for small shrimp than for large shrimp indicating a higher metabolic rate per unit body mass in small shrimp. Wickins (1976) recalculated the specific oxygen consumption values of six species of penaeid prawn (from Egusa, 1961; Kadur, 1962; Kutty, 1969; Kutty et al., 1971 and Venkataramiah et al., 1974) and found mean oxygen consumption to vary from 0.34 mgO, g-’ hr-’ in 1 I g Penaeus japonicus at 23°C to 1.88 mg0, g-’ hr-’ in small 1-2 g Penaeus nztecus at 31°C (Table 6). Silva and Regnauld (1980) measured values of 0.272-0.452 mg0, g-’ hr-’ in Palaemon .verratus, while Hagerman and Weber ( 1981) obtained respira-

Species ___~__

(mg 0, g

~

Penaeus indicus

i I

01 0.1 0.1

MerupenaPw

I

0.5

nlOn0fCTO.Y

3.5 2.70-4.96 1.50-5.31 1.1111.59 2.45-4.3 I I .34-2.29 0.82-4.25 1.52-2.14 0.90-2.04 I 0.61.-14.43 I 2.7 17.4 3.1 (2.4-3.7) 5.5 (4.6-6.2) i ll.R(l3.4-12.6) 16.1 (14.X-18.2) /

I

Penaeus a:lrrus

Penueus in&us Penaerrs semisulcaru.~

Prtmetcs japoni~u\ *Recalculated

by author.

’ hr

tory rates in the same range for ~aia~~~~ a&pews, namely 0.143-0.429 mgOz g-’ hr-‘. Cook and Achituv (in press) investigating the effect of thermal pollution on Palaemon paccjks in the western Cape obtained “basal” respiratory values (recalculated) of 0.4.55-0.899mgOz g-’ hr-‘, and “active” values of 0.761-1.783mgOz g-l hr-’ for 371 mg fresh mass shrimp from 15 to 25°C. This “basal” respiration, which presumably refers to routine oxygen consumption, was approximately double that obtained here using the predictive equation (6). However, they used a closed bottle system which could explain the higher calculated values. The values obtained for P. pacifirus in Table 5 compare well with general shrimp values. Comparisons between slopes and elevations of the same temperatures for different seasons yielded no significant differences showing that no seasonal acclimatization occurs in P. ~u~~~~z~.s.Du Preez (1983)

‘1 for six species of penaeid prawn (from Wtckins.

Mean Oz Live mass (g) consumption I .06 0.97 0.87 1.29 0.941.43 I.18

1.25 i .63 1.60 1.12 1.37

1.65 1.88 0.43-1.13 0.70 0.35 0.58 0.56 0.34 0.41

'

mgO,g-‘hr

31.2 1x2. I 1125.5 il.’ iX2.I 1115.5 ii.2 IX:! I II’55 31.2 1x2. I II?,5 31.2 1x2.1 1175.5 31.1 1x2.1 1125.5 31.2 1x1. I Il’5.S ?I.? 1X2.1 1125.5

w/s

Table 6. The oxygen consumption

pgO,hr-’

T(Y)

S%

25 -28 25-28 25-28 29 30~32 26

7

26 26 26 31 31 31 31 2X.2 30 30 23 23 23 23

17.0 25.5 34.0 8.5 17.0 25.5 34.0 14.5 36.5 36.5 28 28 2X 2X

1976)

Reference Kuttyel

u(.. (1971)

Subrahmanyam Kutty (1969) Kutty (1969) Egusa (1961)

(1962)*

Oxygen

consumption in

investigating respiration in the crab Ovalipes punctatus from the same geographic area and similar temperature range as this study also found no significant differences in elevation between the same temperatures during summer and winter. However, he encountered counter-clockwise slope rotation during winter suggesting a certain degree of metabolic acclimation. Dye (1980) investigating aspects of the respiration of the beach mysid Gastrosaccus psammodytes found no significant differences between winter and summer acclimated males or females but similar to Du Preez (1983) found evidence of a small counterclockwise slope rotation during winter. but only in the case of gravid females. P. pa&us could thus be another example of non-acclimation as was found by Brown et al. (1978) for two species of BuNia. The Q,, values for P. pacjficus varied with temperature but not with mass or season (Table 4), ranging from 2.981 at 30°C to 4.695 at 15°C. This is in accordance with the pattern of higher Q,, values at low temperatures than at high temperatures for Crustacea in general (Wolvekamp and Waterman, 1960). Hart (1980) estimated an average Q,, of 2.13 for Caridina nilotica (2&3O”C) while Cockcroft (1983) obtained Q,,, values of 2.563.158 for the penaeid Mucropetasma qfricanus (15-35’C). Winter acclimated G. psammodJ,tes were found to have higher Q,O values than for summer (Dye, 1980) while Du Preez (1983) found no significant differences between winter and summer Q,, values. A marked increase in oxygen consumption was exhibited by moulting shrimp (Fig. 1). This phenomenon has been well documented in Crustacea which shows that ecdysis is an extremely active metabolic process. Halcrow and Boyd (1967). Buesa (1979). Hart (1980) Alcaraz and Sarda (198 1) and Cockcroft (1983) have all found elevated oxygen consumption rates associated with ecdysis in Gammarus oceanicus, Pam&us cus and

argus. Caridina nilotica. Macropetasma afiicanus

Nephrops

norvegi-

respectively. Kulkarni (1979) and Silva and Regnault (1980) showed oxygen consumption levels to be lowest during intermoult (C) increasing up to premouh (D2-D2) and postmoult (A) for both Emerita holthuisi and Palaemon serratus. respectively. Oxygen consumption varies with animal activity (McFarland and Pickens. 1965). In this study routine metabolism (mean over 24 hr) was measured during which time the shrimp were relatively quiescent. Active metabolism was not measured. but in Caridina fkrnandoi it was 1.1-1.4 times higher than routine metabolism (Wychffe and Job. 1977). Cook and Achituv (in press) measured 1.1882.18 times increases from “basal” to active metabolism in Palaemon pacijicus, while Du Preez (1984) assumed a threefold increase in mean oxygen consumption in the field for Ovalipes punctatus. Estimates for the metabolic requirements of P. paczjkus are probably conservative and would therefore need to be increased by a factor > I in order to better relate to natural conditions when constructing an energy budget for this shrimp. Acknowledgemants~ This research was funded by the South African Council for Scientific and Industrial Research (SANCOR. estuaries programme) and the Department of Environmental Affairs. My thanks are forwarded to I. Davidson and W. Strydom for technical assistance, to

Palaemon

77

Professor A. Dye for discussion and appraisal of the manuscript and Mrs. P. Hawks for preparing the typescript. REFERENCES Alcaraz

M. and Sarda

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tionship

F. (1981) Oxygen consumption by (L) (Crustacea : Decapoda) in relawith its moulting stage. J. exp. mar. Biol. Eco/. norvegicus

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Ser.)

2, 133%141.

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Physiol. 61Ai

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Buesa R. J. (1979) Oxygen consumption of two tropical spiny lobsters Parzulirus argus (Latrielle) and P. guftaru.7 (Latrielle) (Decapoda, Palinuridae). Crustaceana 36, 999104. Cockcroft A. C. (1983) Aspects of the biology of the swimming prawn Macropetasma qfiicanus (Balss). M.Sc. thesis, Univ. Port Elizabeth. South Africa. Cook P. A. and Achituv Y. (1984) The influence of temperature variations and thermal pollution on various aspects of the biology of the prawn Palaemon pacificus (Stimps), in press. Du Preez H. H. (1983) The effects of temperature, season and activity on the respiration of the three spot swimming crab. Ovalipes puncta~us. Cony. Biochem. Physiol. 75A, 353-362. Du Preez H. H. (1984) Consumption. assimilation and energy balance in the three-spot swimming crab Owlipes puncfatus (De Haan) (Crustacea: Brachyura) In Developments

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(Edited by McLachlan A. and Erasmus T.), Vol. 19, pp. 637-642. W. Junk, The Hague. Dye A. H. (1979) The effect of acute and long term temperature changes on the respiration of two sand-dwelling bivalves. Contp. Biochmr. Physiol. 63A, 405409. Dye A. H. (1980) Aspects of the respiratory physiology of Gastroscrccus psamnzo&tes Tattersall (Crustacea : Mysidacea). Camp. Biochem. Physiol. 6SA. l87- 19 1. Dye A. H. and McGwynne L. (1980) The effect of temperature and season on the respiratory rates of three psammolittoral gastropods. Camp, Biochem. Physiol. 66A, 107-111.

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