Hatching Time in Spherical, Pelagic, Marine Fish Eggs in Response to Temperature and Egg Size8

Environmental Biology of Fishes Vol. 22, No.4, pp. 261-271, 1988 © K1uwer Academic Publishers, Dordrecht.

Hatching time in spherical, pelagic, marine fish eggs in response to temperature and egg size8

Daniel Pauly & Roger S.V. Pullin International Center for Living Aquatic Resources Management (ICLARM), MC PO Box 1501, Makati, Metro Manila, Philippines

Received 29.8.1986 Accepted 4.U.1987

Key words: Teleosts, Early life history, Embryology, Environmental physiology

Synopsis

The relationship between egg diameter (, mm), temperature (T, ° C) and egg development time to hatching (D, in days) was established for approximately spherical, pelagic marine fish eggs as 10g100 = 7.10 + 0.608 10g)0 - 4.09 log)o (T+ 26), which explains 82% of the variance of a data set of 140 cases, covering 84 species of teleost fishes, for temperatures from 2.8 to 29.5°C and eggs of 0.6 to 3.4mm diameter. From this relationship it appears that temperature has 4.7 times as much effect on development time as egg diameter when all variables are expressed in standard deviation units. A discussion of these and related factors is given.

Introduction temperature fluctuations) on hatching time can be used in the context of commercial farming and A strong, inverse relationship between the time stocking ventures (e.g., Hayes 1949, Alderdice & needed for fish to develop within the egg envelope Velsen 1978, Crisp 1981). and ambient water temperature has been reported The effects of factors other than temperature, as early as the 18908 (Dannevig 1895) and this rela­ such as salinity and dissolved gases, on the devel­ tionship has been studied quantitatively by a num­ opment offish eggs have been less investigated and ber of authors (Apstein 1909, Johansen & Krogh again most studies refer to freshwater or brackish­ 1914, Blaxter 1956, Ignatyeva 1974, Ryland & Ni­ water species (e.g., Alderdice et al. 1958 and stud­ chols 1975, Russell 1976, Martin & Drewery 1978, ies reviewed by Hempel 1979). Breder & Rosen Hoestlandt & Devienne 1980, Jones et al. 1978, (1966) have also reviewed scattered reports on the Thompson et al. 1981). Blaxter (1969) states that effects of various physico-chemical factors. 'the time to hatching is both a specifically and envi­ In this study, we have confined our attention to ronmentally controlled characterwith ternperature the effects of temperature and egg size (diameter) and oxygen supply exerting a consiqerable effect'. on the development time to hatching of approxi­ Most studies have been conducted with freshwater mately spherical, pelagic, marine eggs. It is appar­ species, particularly salmonids, in which a precise ent that large eggs develop more slowly than small knowledge of the influence of temperature (and eggs, other factors being equal (see data compiled in Table 1). Our purpose is not to demonstrate •ICLARM Contribution No. 308. these phenomena, but rather to quantify them to 262

Table 1. Data set for developmeot time to hatching (D, days), egg diameter (41, mm) and water temperature (T, 0c) for spherical, pelagic, marine fish eggs (for criteria of data selection, sec text).

Family Species D (days) 41 (mm) Temp (0C) Reference

Clupeidae Etrumeus teres 1.50 1.33 20.5 Jones et al. (1978) 2.00 1.33 24.0 Jones et al. (1978) 2.08 1.33 21.5 Jones et al. (1978) 5.63 1.33 11.0 Jones et al. (1978) Brevoortia tyrannus 2.00 1.65 17.5 Jones et al. (1978) Sprat/us sprattus 11.50 0.99 4.3 Thomson et al. (1981) 9.33 0.99 5.2 Thomson et al. (1981) 7.83 0.99 6.0 Thomson et al. (1981) 6.75 0.99 7.0 Thomson et al. (1981) 5.79 0.99 8.0 Thomson et al. (1981) 5.17 0.99 8.9 Thomson et al. (1981) 4.43 0.99 9.7 Thomson et al. (1981) 4.06 0.99 10.6 Thomson et al. (1981) 3.65 0.99 11.4 Thomson et al. (1981) 3.36 0.99 12.2 Thomson et al. (1981) 2.95 0.99 13.2 Thomson et al. (1981) 2.77 0.99 13.7 Thomson et al. (1981) 2.48 0.99 14.8 Thomson et aI. (1981) 2.32 0.99 15.6 Thomson et al. (1981) 2.17 0.99 16.5 Thomson et al. (1981) 2.13 0.99 17.4 Thomson et al. (1981) 1.88 0.99 18.4 Thomson et aI. (1981) 1.80 0.99 19.1 Thomson et al. (1981) 1.76 0.99 20.0 Thomson et al. (1981) Sartlinops ocel1llla 3.70 1.82 11.0 King (1977) 1.75 1.82 16.0 King (1977) Dussumeria luuselti 1.50 1.50 28.5 Delsman (1972) Chanidae Chanos chanos 1.06 1.16 29.5 Liao et al. (1979) 1.19 1.13 28.2 Chaudhuri et al. (1978) 1.04 1.13 28.2 Chaudhuri et aI. (1978) Gadidae Gadus morhua 10.50 1.50 8.3 Breder & Rosen (1966) Pollachius virens 9.00 1.15 9.4 Breder & Rosen (1966) Melanogrammus aegleflllUS 15.00 1.50 2.8 Breder & Rosen (1966) 13.00 1.50 5.0 Breder & Rosen (1966) Micromesistius poutassou 11.50 1.16 8.0 Russell (1976) 4.00 1.16 10.5 Russell (1976) Urophycis regius 2.38 0.71 22.5 Hardy (1978) Urophycis chuss 1.25 0.74 21.1 Hardy (1978) Enchelyopus cymbrius 5.40 0.96 13.0 Hardy (1978) 4.50 0.96 15.0 Hardy (1978) Merluccidae Merluccius merluccius 10.00 1.10 9.1 Breder & Rosen (1966) Merluccius bilinearis 2.00 0.88 21.0 Hardy (1978) Merluccius aJbidus 7.00 1.10 9.8 Hardy (1978) F1stulariidae F/Stularia se"alil 4.00 1.60 28.5 Delsman (1972) Triglidae EU/rigliJ gumardui 5.00 1.41 15.0 Russell (1976) Lepido/rigliJ japonica 2.29 1.30 20.0 Breder & Rosen (1966) Chelidonichthys kunw 7.00 1.70 9.0 Breder & Rosen (1966) Priono/us carolinus 2.50 LOS 22.0 Fritzsche (1978) 3.71 LOS 20.5 Fritzsche (1978) Platycephalidae PlatycephaJus indicus 1.00 0.90 25.0 Breder & Rosen (1966) Centropomidae La/es caicarifer 0.70 0.87 27.5 Wongsomnuk & Manevonk (1973) 263

Table 1. Continued.

Family Species D (days) 41 (mm) Temp (0C) Reference

Serranidae Epinephelus tauvina 1.27 0.77 28.5 Hussain et al. (1975) 1.00 0.90 27.0 Chen et aI. (1977) Dicentrarchus labrax 4.67 1.18 13.0 Bamabe (1976) 2.50 1.35 15.0 Russell (1976) 2.29 1.35 17.0 Russell (1976) Centropristes striatus 5.00 1.00 10.0 Breder & Rosen (1966) 5.00 0.95 10.0 Hardy (1978) 5.00 0.95 15.0 Hardy (1978) 3.13 0.95 15.0 Hardy (1978) 3.13 0.95 16.0 Hardy (1978) 1.58 0.95 23.0 Hardy (1978) Laleolabrax japonicus 4.50 lAO 13.0 Breder & Rosen (1966) Pomatomidae POTrUltomus saitatrix 1.96 1.00 20.0 Hardy (1978) Sillaginidae Sillago sihama '0.83 0.66 24.5 Breder & RoseD (1966) Carangidae Caranx crumenophthalmus 0.63 0.78 28.0 Delsman (1972) Decapterus kurra 0.50 0.70 28.5 Delsman (1972) Decapterus macrosoma 0.38 0.62 28.5 Delsman (1972) Serioia quinqueratliata 2.08 1.00 21.0 Kuronuma & Fukusho (1984) Coryphaenidae Coryphaena hippurus 2.00 1.40 24.5 Johnson (1978) Leiognathidae Leiognathus nuchalis 1.56 0.60 23.0 Breder & Rosen (1966) Lutjanidae Lutjanus iulsmira 0.75 0.82 26.4 Suzuki & Hioki (1979) Lethrinidiae Lethrinus nl!TrUltacanthus 1.63 0.81 20.4 Breder & Rosen (1966) Sparidae Stenotomus chrysops 1.67 1.00 22.0 Johnson (1978) 1.67 0.90 22.2 Breder & Rosen (1966) Archosargus probatocephalus 1.67 0.80 25.5 Breder & Rosen (1966) Pagrosomus auratus 1.88 1.00 18.0 Breder & Rosen (1966) Mylio TrUlcrocephaius 2.50 1.04 19.3 Fukhara (1977) Acanthopagrus cuvieri 1.66 0.81 21.0 Hussain et aI. (1981) Nemipteridae Nemipterus variegatus 1.17 0.68 24.0 Breder & Rosen (1966) Sciaenidae Bairdiella chrysoura 0.75 0.76 27.0 Johnson (1978) 1.88 0.76 20.0 Johnson (1978) Poganias chromis 1.00 0.92 20.0 Johnson (1978) MenJicirrhus saxatilis 2.00 0.82 20.5 Johnson (1978) 2.08 0.84 20.0 Breder & Rosen (1966) Nibea argentata 0.92 0.76 23.0 Breder & Rosen (1966) Lagodon rhomboitks 2.00 1.02 18.0 Cardeilac (1976) Oplegnathidae Oplegnathus fasciatus 1.50 0.91 21.0 Breder & Rosen (1966) Ephippidae Chaetodipterus faber 1.00 1.00 27.0 Johnson (1978) Mugilidae Mugil cephalus 1.54 0.93 24.0 Kuo et ai. (1973) 2.04 0.93 22.0 Kuo et ai. (1973) Mugil macrolepis 0.96 0.67 27.5 Sebastian & Nair (1975) Sphyraenidae Sphyraena pinguis 1.13 0.76 23.7 Breder & Rosen (1966) Labridae Tautogalabrus atlsperus 1.67 0.85 21.5 Fritzsche (1978) Tautoga onitis 1.81 1.00 21.1 Breder & Rosen (1966) Thalassoma cupido 1.50 0.48 23.3 Breder & Rosen (1966) Scaridae Calotomus japonicus 1.00 0.66 25.0 Breder & Rosen (1966) Trachinidae Trachinus draco 4.50 1.04 16.8 Russell (1976) Acanth 'd t: Acanthurus triostegus 1.08 0.67 24.0 Breder & Rosen (1966) 5iganidae Siganus argenteus 1.04 0.65 26.5 Burgan & Zseleczky (n.d.) Gempylid= Thyrites a/un 2.08 0.99 18.5 Breder & Rosen (1966) Trichiundae Trichiurus sp. 2.00 2.42 28.5 Delsman (1972) Scombrid e Scomber scomber 7.38 1.19 7.4 Russell (1976) 2.06 1.19 21.0 Russell (1976) 264

Table]. Continued.

Family Species D (days) $ (mm) Temp (0C) Reference

Scomber japonicus 2.04 1.05 19.5 Fritzsche (1978) 2.08 1.05 20.0 Fritzsche (1978) Thunnus albacores 1.85 0.96 18.7 Harada et al. (1980) 1.40 0.96 24.4 Harada et aI. (1980) 1.34 0.96 30.1 Harada et a!. (1980) Thunnus obesus 0.88 1.05 28.8 Fritzsche (1978) KalSuwonus pe/amis 1.10 1.00 26.7 Inoue et al. (1974) Scomberomorus macu/atus 1.04 1.03 25.0 Breder & Rosen (1966) 0.65 1.00 29.0 Fritzsche (1978) 1.02 1.00 25.5 Fritzsche (1978) Nomeidae Seriolella punctata 6.08 1.14 11.5 Grimes & Robertson (1981) Stromateidae Peprilus triacanthus 3.00 0.75 14.6 Martin & Drewery (1978) Bothidae Paralichthys dentatus 2.33 1.02 22.9 Martin & Drewery (1978) 3.06 1.02 17.5 Martin & Drewery (1978) 5.92 1.02 9.1 Martin & Drewery (1978) Scophthalmus maeotkus 7.00 1.10 11.5 Martin & Drewery (1978) 5.42 1.10 13.5 Martin & Drewery (1978) 5.29 1.10 14.0 Martin & Drewery (1978) 5.17 1.10 14.2 Martin & Drewery (1978) 5.00 1.10 15.0 Martin & Drewery (1978) 4.71 1.10 16.3 Martin & Drewery (1978) 3.00 1.10 17.7 Martin & Drewery (1978) Scophthalmus maximus 9.50 1.06 10.0 Russell (1976) 7.00 1.06 12.0 Russell (1976) 5.00 1.06 14.5 Russell (1976) Zeugopterus punctatus 3.00 1.00 14.5 Russell (1976) Pleuroneetidae Limanda limanda 7.00 0.93 9.0 Russell (1976) 12.00 0.93 7.0 Russell (1976) 3.00 0.93 10.0 Russell (1976) Microstomus kilt 5.50 1.29 15.3 Russell (1976) 8.80 1.29 8.8 Russell (1976) 6.00 1.29 6.0 Russell (1976) Glyptocephalus cynoglossus 8.00 1.16 8.6 Russell (1976) Hippoglossus hippoglossus 16.00 3.40 6.0 Russell (1976) Kareius bicoloratus 9.00 1.03 5.0 Yusa (1979) Ostraciidae Lactophrys quadricornis 2.00 1.46 27.3 Breder & Rosen (1966)

derive a relationship for predicting development dard methods of fishery biology, such as egg sur­ time, given temperature and egg size in spherical, veys, for the determination of spawning stock bio­ pelagic, marine fish eggs, irrespective oftheir taxo­ mass (Saville 1977). nomic affinities. A secondary aim of this contribution is to dem­ onstrate that the small egg sizes of most tropical Material marine fishes, combined with the high temperature of their habitat, usually cause egg development Table 1 presents the basic data used for this study. time to be extremely short, hence precluding (or The 140 records (originally 141, see further) pre­ making very difficult) the application ofsome stan- sented there were assembled from the literature, using the following selection criteria: (i) eggs nored. Also, in view of the scattered nature of our should have developed in full-strength seawater data, no attempt was made to model responses by (which excludes the effect ofsalinity from our con­ taxonomic grouping. siderations); (ii) eggs should have a spherical shape (which excludes engraulid eggs, among others); (iii) eggs should lack tendrils or other protuber­ Models and methods ances (such as occur in flying fish eggs (Breder & Rosen 1966) and cause the eggs to stick together The 141 data sets initially available were fitted with and to floating objects) or significant chorionic a multiple linear regression model of the form sculpturing (Robertson 1981); (iv) eggs should be pelagic (as opposed to demersal, e.g. herring eggs); (1) and (v) each published value used for development time (in days) should be matched with a co-publish­ where D is the development time in days (Le. time ed orwell established value for the egg diameter (in to hatching), ct> the egg diameter in mm and T the

mm) and for the temperature prevailing during the water temperature in 0 C. This exploratory model incubation period. was used for two purposes: (i) to identify outliers Criteria (i) to (iv) were meant to ensure as much (Le. values producing very high residuals), and (ii) I uniformity as possible within the data used. Crite­ to provide a baseline value of the goodness of fit rion (v) was, in some cases fulfilled by matching a estimator (R2), as required for assessing whether hatching time/temperature record with an egg size more complex models do indeed provide a better value from another source, usually an identifica­ fit. tion guide. In such cases, Table 1 gives only the Only one clear outlier (see Fig. 1 and text below) source of the development time/temperature rec­ was identified in this first pass. Yet, still, it provid­ ord. In some other cases, criterion (v) was fulfilled ed a value of (corrected) R2 = 0.821. by matching a development time/size record with a A second multiple regression model was run temperature taken from another source, usually an then. It was based on the l40-row data set in Table appropriate mean field measurement from the ge­ 1, and included a squared term (T2) as an additional ographical range. In such cases, Table 1 gives only term which, if significant, would suggest that sim­ the source of the development time/size record. ply taking logarithms is not sufficient to linearize Whenever necessary, ranges (of development the temperature term in equation (1). We found time, size and/or temperature) were replaced by that such squared term was indeed significant and midrange values. We expect some of the unex­ hence used for our final analysis a model of the plained variance in the models presented below to form stem from some of the methods by which the data were compiled. Altogether, 84 species, representing 35 families of teleost fishes are included here, with egg dia­ The value of a was varied in steps of 10 C until the meters ranging from 0.6 to 3.4 mm (but mostly highest possible value of R2 was achieved. Note within 0.75-1.50mm), temperatures ranging from that, to obtain a dimensionally correct interpreta­ 2.8 to 29SC and development times ranging from tion, 'a' must be understood as having the dimen­ about 9 hours to 12 days. This data set represents a sion time; bl> the dimension timellength and b2, the good cross-section of pelagic marine eggs since ac­ dimension time/temperature. All computations cording to Ahlstrom & Moser (1980) over 40% of were performed on an Osborne Executive I Micro­ such eggs are <1.0mm diameter, 30% are between computer using the STATPAK Multifunction Sta­ 1.0 and 1.5 mm, 15% are between 1.5 and 2.0 mm, tistical Library of Northwest Analytical Inc., Port­ and about 14% are >2.0mm. The presence, ab­ land, Oregon. sence and/or configuration of oil globules were ig- 266

15

• II) ->- 0 • -0 • Q) E 10 - • -c • -Q) • E 0- • 0 • Q) > • •• Q) - 0 • • • • e 5 •• •, • • e• • • • -. .., e -. - -.• •• oLllller ment oned In te . @ • oO=-----J'------::O::----I.---::2~O----'----:3.LO-----I Tem,pemune (oCl

Fig. 1. Development time to hatching ofthe near spherical marine fish eggs documented in Table 1, ploued against water temperature. This plot does not take account of different egg sizes (but see lext).

Results 'about 1mm in diameter, and [which] hatch in about 12 hours at a temperature ofabout 12° C' (see Table 2 presents the statistics of the multiple re­ Fig. 1). This record was deleted from all subse­ gression used as a first pass model; it has the form: quent analysis. Running a multiple regression with a squared

logloD = 2.09 + O.619Iog lO$ - 1.381l0gloT. (3) term produced the equation

This model produced an extremely large residual 10gIOD = 0.886 + 0.608IoglO$ +

for the data on Prolalilus jugularis (Branchiostegi­ O.8831og1OT- l.021log10'f2, (4) dae) originally generated by Fischer (1958) and reported by Breder & Rosen (1966, p. 455) as eggs with R2 = 0.850, but with F= 257.585, which is 267 lower than the F-value for equation (3) (see Table all linearization of the model was achieved. 2). Expressing the variables in equation (4) in stan­ The best fit for a model such as (2) was obtained dard deviation (s.d.) units led to: with a value of Q = 26° C, Le. 0' = 0.196 cI>' - 0.992 T' (6) 108100 = 7.10 + 0.60810glocI>­ 4.0910g10 (T+ 26). (5) in which 0', cI»' and T' are 10g100, 10glOcI> and loglo (T + 26) expressed in s.d. units, while 0.196 and The statistics of this model are given in Table 3. 0.922 are 'path coefficients', Le. standardized par­ Figures 2 and 3 give plots ofthe residuals on 10glocI> tial slopes, which express in comparable units the and loglo (T+ 26), respectively, while Figure 4 effect on a given process of variables that were gives a plot ofthe actual on the predicted values. It originally expressed in different units (Li 1975). It will be noted that the residuals in Figures 2 and 3 will be noted that temperature has, in the data set are randomly distributed. Figure 4 shows that over- used here, an effect 4.7 times (= 0.922/0.196) larger than cI>.

Table 2. Statisticsofmultiple regTession between loglo development time to hatching, IOg10 egg diameter and 10g10 temperature (see text, equation 3).

Tenn Value of CQc:Ificlcnt Standard error T-SIotis[ic Partial colTdal ion

EO 2.08'.507 0.079616 26.1B9 Bl 0.6]927 O. 11 8M2 5.228 0.4019 B2 IJ807J 0.064536 -21.396 - 0,8773

Sum of squares Degrees of freedom Mean squares

Due to regression 13.7069 2 6.85345 About regression 2.9459 137 0.02150 Total 16.6528 139 0.11981 R2 0.8231 Corrected R2 0.8205 F-Test 11 318.7190 Std. error of regr. 0.1466

Table J. Sta.istics of multiple regression between loglo development time to hatchings, 10g10 egg diameter and 10g10 temperature index (T + 26) (equation 5; see Table 2 and text).

Term Value of coefficient Standard error T-statistic Panial correlation

BO 7.]0326 0.28152 2:5.1425 Bl 0.60166 ''-1090.3 S.Si33 0.'1299 B2 4.01rl66 0.17220 23.7],'13 - 0.8969

Sum of squares Degrees of freedom Mean squares ! I Due to regression 14.1513 2 7.07563 About regressi~n 2.5016 137 0.01826 Total 16.6528 139 0.11980

R2 0.8498 Corrected R2 0.8476 F·Tcst .Il 387.5010 Std. error of regr. 0.1351 268

0.50

0.25 • • • • .. • •• •• .. • • • • •• • • •• • •• • • ••• • •• • • •• ••• ••• • ••••• o ---- • ... ------.------en .------.-.- ...... o •• • • •• •• • • ••• :::J ..... - ...... '0 • •• • • • ••• en Q) • • a:: -0.25 • • • • -0.50 1.5 1.6 1.7 loglo Temperature index (OC+26)

Fig. 2. Plot of rC5iduals ofequation (5, and5CC text) against loglo temperaturc index (OC + 26). Small dols represent single values; large dOIs, two or more overlapping values.

0.50

0.25 • • •••• en • ...... ·c- • • ••.. . :::J . - . ~ 0 ------.---.. ------'--- o .. ---.-'.---. • • ••• • •• • en ...... o •••• • • • :::J • •• '0 'iii - 0.25 • Q) a::

-0.50_0.3 -0:'2 -0.1 0 0.1 0.2 0.3 0.4 0.5 loglo Egg diameter (mm)

Fig. 3. Plot ofresiduals ofequation (5, and sec text) against loglo egg diameter (mm), Small dols represent single values; large dOIs, two or more overlapping values. 269

• .2 ~ "."", 0) E ----,- t,· 1.0 •• r'.'• c " -0) .. " ~.'. .. E 0.8 • ..• -fI".• •• .. Q. 0_ . .. /~ . ~~0.6 ••• ,;1 .. 0)0 .,." '0'0 . ...."It.- "0-0.4 ...... 0) _ -U . .,...... ~ .. '0 0.2 0) . .M...... " " ... . a. .... ,N • Q 0 CI . ., ,'.. .. o _c.e_~·_'L-.,o:.::'_...L__L __L._.....L__-..L__-L__...1- -0.4 -0.2 0 0.2 0.4 0.6 0.8 1.2 1.2 loglo Actual development time (days)

Fig. 4. Plot of predicted against actual egg development time to hatching in days (see equation 5 and text). Small dots represent single values; large dots, two or more overlapping values.

Discussion tant (see below). The best model proposed here (equation 5) should nevertheless be of help in esti­ The key problem associated with the data set pre­ mating the development time of typical marine fish sented here is that it is very heterogeneous. Differ­ eggs under conditions where high precision is not ences within the data set could be from three sourc­ required, as in ecosystem simulations. Moreover, es: such models are a step toward a compendium of (i) those caused by different methods andcriteria basic early life history data constituted around a for determining development time, which should carefully selected format, as suggested by Alder­ ideally refer to the time when 50% ofembryos have dice (1985). hatched; Equation (5) implies that from a small egg of say (ii) true taxonomic differences in species-specific 0.6mm diameter developing in the Java Sea (at development rate; 28,50 C) an embryo will hatch within about 18 h (iii) random effects, associated with our method after fertilization, and hence will spend only a few of data compilation. hours in each of the stages into which one might Multiple regression models of the form presented wish to split the development process. This will here can account only for the variance caused by severely hamper the use of egg surveys to estimate (iii), and will in fact tend to compound this with the the spawning stock biomass ofmany tropical fishes effects of (i) and (ii). We cannot, for this reason, as used in cold water stocks (Saville 1977), because resolve on the basis of the data set in Table 1, the this method implies that 'an estimate of number of question of whether true taxonomic differences in eggs produced [is] obtained by sampling eggs of development rate occur after standardization for known age at a series ofpoints in time and location egg diameter and temperature. However, we sus­ and then integrating over area and time to get the pect that such taxonomic differences are not imper- total production'. 270

Finally, we pose the question whether models daurade Sparus aurala (L.). Etud. Rev., Cons. Gen. Peches such as derived herein .from a large and hetero­ Mediterr. 55: 63-116. geneous data set, have any implications for life Blaxter, J.H.S. 1956. Herring rearing. II. The effect of temper­ history strategy. According to available evidence ature and other factors on development. Mar. Res. Scotl. 5: 1-19. reviewed by Balon (1984), hatching time is of little Blaxter, J.H.S. 1969. Development: eggs and larvae. pp. 177­ survival value, at least in comparison with the later 252. In: W.S. Hoar & D.J. Randall (ed.) Fish Physiology, threshold of transition to exogenous feeding. Vol. 3, Academic Press, New York. Our data show that any egg of a given size is Breder, C.M., Jr. & D.E. Rosen. 1966. Modes ofreproduction in fishes. T.F.H. Publications, Jersey City. 941 pp. 'driven' by the prevailing temperature to the point Burgan, B.G. & K.A. Zseleczky. n.d. Induced spawning and ofhatching, since like any othersystem it is subject early development of the rabbitfish, Siganus argenteus (Quoy to the laws of thermodynamics and since it is be­ and Guimard) in the Philippines. Contribution from the Ma­ yond all doubt completely poikilothermic. In other rine Laboratory, Silliman University, Dumaguete City, Ne­ words, such eggs cannot help but hatch 'at the gros Oriental, Philippines. 9 p. (Mimeo). Cardeilac, P.T. 1976. Induced maturation and development of mercy' of the prevailing temperature. This may pinfish eggs. Aquaculture 8: 389-393. seem obvious, but it is further useful evidence Chaudhuri, H., J.V. Juario, H. Primavera, R. Samson & R. against hatching as a decisive event with respect to Mateo. 1978. Observations on artificial fertilization of eggs survival value. The scope for regulation oflocomo­ and the embryonic and larval development of milkfish, Cha­ tory activity and other processes leading up to ex­ nos chanos (Forskal). 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