FISHERIES
Fish reproductive-energy output
increases disproportionately
with body size
Diego R. Barneche,1*† D. Ross Robertson,2 Craig R. White,1 Dustin J. Marshall1
Body size determines total reproductive-energy output. Most theories assume
reproductive output is a fixed proportion of size, with respect to mass, but formal
macroecological tests are lacking. Management based on that assumption risks
underestimating the contribution of larger mothers to replenishment, hindering
sustainable harvesting. We test this assumption in marine fishes with a phylogenetically
controlled meta-analysis of the intraspecific mass scaling of reproductive-energy output.
We show that larger mothers reproduce disproportionately more than smaller mothers
in not only fecundity but also total reproductive energy. Our results reset much of the
theory on how reproduction scales with size and suggest that larger mothers contribute
disproportionately to population replenishment. Global change and overharvesting cause
fish sizes to decline; our results provide quantitative estimates of how these declines
affect fisheries and ecosystem-level productivity.
H
ow does reproductive output scale with
body size? Despite the august history of
this question (1, 2), theoreticians, fisheries
scientists, and field biologists still disagree
about the answer (tables S1 and S2). Re-
solving this uncertainty is essential for under-
standing the forces driving the evolution of body
size in general and for managing fish stocks sus-
tainably. Most life-history models, mechanistic
theories of growth, and fisheries models assume
that reproductive output scales isometrically with
femalemass (table S1)—that is, for every increase
in female somatic mass, there is a constant pro-
portional increase in reproductive output (Fig. 1A,
dashed line). Under such a model, the reproduc-
tive output of one 2-kg fish is equal to that of two
1-kg fish.
In contrast to most theoreticians and fisheries
models, field biologists have repeatedly suggested
that fish fecundity may increase disproportion-
ately with body mass within species (that is, it
scales hyperallometrically; table S2) (3, 4). Thus,
two 1-kg fish would have less reproductive out-
put than a single 2-kg fish (Fig. 1). This discrep-
ancy between theory and empiricism takes on
particular importance because reproductive out-
put drives the replenishment of fisheries (3).
Energy investment into individual offspring may
also change with female size, such that larger
mothers produce larger offspring, which survive
better (4, 5). Many life-history models make ex-
plicit assumptions about the energy devoted to
reproductive tissue (table S1). Therefore, under-
standing how reproductive output scales with
female size requires not only fecundity estimates
but also estimates of how egg size and egg energy
scale with female size. Such an understanding
would (i) improve the management of exploited
populations and protected areas by better esti-
mating the relationship between standing bio-
mass and egg production (3, 4, 6) and (ii) allow
direct tests of energy-budget models that char-
acterize changes in reproductive allocation over
ontogeny (7).
Here we perform a meta-analysis of intraspe-
cific mass scaling of total reproductive-energy
output (fecundity × egg volume × egg energy),
with controls for phylogenetic nonindependence,
for 342 species of marine fishes from 15 orders
(8). We collected data on how egg energy content
scales with egg volume within and among a sub-
set of species (n = 1366 clutches from 126 species).
We estimated scaling of both egg characteristics
and fecundity because these traits could trade off
against each other, resulting in no net relation-
ship between maternal size and reproductive
output. For example, if larger mothers produce
more, but smaller, offspring, net reproductive
output might not change with maternal size.
Fish fecundity varied from 11 to 57,600,000
eggs per clutch, spanning about seven orders of
magnitude. Fish fecundity scaled hyperallomet-
rically (a power-function relationship with an
exponent greater than 1) with body mass (8)
(Fig. 2A). In this analysis, 140 (79.1%) of the 177
species presented hyperallometric mass scaling
of fecundity.
The volume of individual eggs varied from0.01
to 344.8 mm3 and increased hypoallometrically
with female body mass. A 1.2-fold increase in egg
volume would be expected with an increase in
body mass from 0.3 to 1.3 kg (Fig. 2B), hence
strengthening the effect of female mass on total
reproductive output. Considering only the spe-
cies for which we had both fecundity and egg-
volume data (n = 45), we find a hyperallometric
relationship between female mass and total re-
productive volume (that is, fecundity × egg volume)
[average scaling exponent = 1.21; 95% credible
interval (CI) 1.07 to 1.37; table S6] (8).
Energy content of individual eggs varied from
0.07 to 299.41 J. Egg energy scaled hypoallometri-
cally with egg volume (Fig. 2C). That is, larger eggs
have slightly less energy content per unit volume
than smaller eggs but still have a greater energy
content in absolute terms. The combination of
this relationship and the relationship between
female mass and egg volume still yields a posi-
tive, hypoallometric relationship between female
size and per-egg energy content (8).
By combining the posterior distributions of
model parameters from the three models above
(8), we were able to estimate how the average
total reproductive-energy output changes with
body size for marine fish. Total reproductive-
energy output per spawning scales hyperallo-
metrically with female body mass (mean scaling
exponent = 1.29; 95%CI 1.20 to 1.38). An estimate
of mass scaling of fecundity in the commercially
important Atlantic cod (Gadus morhua) illus-
trates these scaling effects. If fecundity scaled
isometrically with mass, then 15 2-kg female
cod would produce the same number of eggs as
1 30-kg female. Instead, we find that a single
30-kg female produces more eggs than ~28 2-kg
females (weighing a total of 56 kg). Further, be-
cause egg volume and egg energy content also
increase with female size, a 30-kg female actually
spawns a batch of eggs with a total energy con-
tent ~37 times as high as that of a batch of eggs
from a single 2-kg female (Fig. 1B); assuming an
isometric relationship would underestimate this
difference by 147%.
Unsurprisingly, substantial variation in
reproductive-energy output exists among spe-
cies, and a small proportion of species (5.0%)
show isometric or hypoallometric scaling (Fig. 3).
Nevertheless, hyperallometric reproductive scal-
ing holds for almost all species (95.0%) and is
consistent for the subset of nine species for which
we had data on all three relationships of interest
(table S7) and for three studies in which all the
data came from the same populations (table S8).
Thus, larger females have disproportionately
higher fecundity and produce offspring of greater
size, content, and possibly quality. Assuming that
larger mothers take part in at least as many
reproductive bouts each reproductive season
as smaller mothers [which seems likely (4)],
these relationships may explain why larger fish
are so important for the replenishment of marine
fish populations (6, 9).
Marine protected areas (MPAs) increase the
size of fish by 28% on average (7). Because the
per-capita reproductive output of fish increases
with size, the potential for MPAs to replenish
populations has been underestimated. For ex-
ample, for thewidow rockfish,Sebastes entomelas,
anMPA could enhance population replenishment
by 60 and 74% for fecundity and reproductive-
energy output, respectively—no such increase
would be predicted if reproductive output was
assumed to be isometric (8).
RESEARCH
Barneche et al., Science 360, 642–645 (2018) 11 May 2018 1 of 3
1Centre for Geometric Biology, School of Biological Sciences,
Monash University, Clayton, VIC 3800, Australia.
2Smithsonian Tropical Research Institute, Balboa, Panama.
*Present address: School of Life and Environmental Sciences, The
University of Sydney, Sydney, NSW 2006, Australia.
†Corresponding author. Email: barnechedr@gmail.com
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Our results also reveal the insidious costs of
global change. Fish sizes are predicted to de-
crease in some instances as temperatures rise
with global warming. For example, a 1.5°C in-
crease in sea surface temperature will decrease
fish lengths by ~15% in the Mediterranean (10).
On the basis of our estimates, such a size de-
crease would incur a 50% per-capita reduction
in fecundity for Atlantic mackerel, Scomber
scombrus (8). Such effects would exacerbate
the impacts of predicted decreases in total fish
biomass associatedwithwarming—warmer oceans
will likely have fewer fish and much lower re-
productive output (11).
Here we show that hyperallometry of
reproductive-energy output is not the exception
but rather the rule for marine fishes, a relation-
ship that fisheriesmodels nowneed to incorporate.
For example, balanced harvesting approaches to
fishing, which are the subject of intense debate
(12, 13), emphasize somatic productivity alone
and assume isometric reproduction. Our results
confirm the suspicion (3) that such assumptions
severely underestimate the importance of larger
females for population replenishment (6, 7, 9).
Worryingly,many exploited species have declined
in size as result of overharvesting, and our find-
ings suggest that such declines will massively re-
duce reproductive output and recruitment (14).
Because our results formally establish a gen-
eral hyperallometric body-mass scaling of repro-
ductive output in marine fishes, they also have
direct implications for fundamental theory.Most
theories of growth and life history assume that
reproductive output scales isometrically with size,
hence underestimating the advantages of grow-
ing larger, although some branches of theory do
anticipate our findings (table S1). However, even
among studies that anticipate hyperallometric
scaling, some assume that energy content of
reproductive tissues stays constant with body
size (table S1); our results contradict this core
assumption for fishes, at least.
Future studies will establish whether our re-
sults are general across groups of organisms other
thanmarine fishes, thus helping determine the
generality of conclusions about the putative evo-
lutionary advantages of large size to mothers. All
Barneche et al., Science 360, 642–645 (2018) 11 May 2018 2 of 3
Female mass (g)
Fe
cu
n
di
ty
 (#
 eg
gs
/fe
ma
le)
1 1 x 102 9.5 x 103
10−1
101
103
105
107
y 2.93 x1.18
95% C.I. :  1.12 1.24
n 7,737
177 spp.
A
Female mass (g)
Eg
g 
vo
lu
m
e 
(m
m3
)
8.8 x 10 1 68 5.2 x 103
0.06
0.14
0.35
0.85
2.08
y 0.15 x 0.14
95% C.I. :  0.06 0.22
n 926
154 spp.
B
Egg volume (mm3)
Eg
g 
en
er
gy
 (J
)
0.07 0.27 1.14 4.8 20.14
0.2
0.5
1.9
6.5
22.7
y 2.15 x0.77
95% C.I. :  0.68 0.86
n 1,366
126 spp.
C
Fig. 2. Scaling relationships of fish reproductive traits. (A) Relationship
between fecundity and female mass. Red circles represent individual clutches.
(B) Mass scaling of egg volume. (C) Volume scaling of egg energy content.
Green triangles represent independent observations [egg volume and egg
energy content in (B) and (C), respectively].The y axes were corrected for
species-specific deviations on the intercept and scaling exponent, which were
obtained using Bayesian phylogenetic hierarchical modeling. Equations in the
top-left corners depict average fixed effects; 95%CI is Bayesian credible interval
for the scaling exponent; n is the total number of observations. Dashed black
lines depict average model fits. Species and family names and the data
summary and contributions for the threemodels [shown in (A), (B), and (C)] (8)
are provided in tables S3 to S5, respectively. Note both axes are logged.
Female mass x 103, M i , (g)
R
ep
ro
du
ct
ive
 o
u
tp
ut
, R
i,
 
(M
J)
A
0
20
40
60
80
0 10 20 30 40
Ri = 67.75 × 10−6 × M i β1
Hyper−allometric
scaling
β1 = 1.33
Isometric
scaling
β1 = 1
(2 kg)
(30 kg)
n = 1
∑ Ri ≈ 61 MJ
∑ M i = 30 kg
n = 37
∑ Ri ≈ 61 MJ
∑ M i = 74 kg
B
Fig. 1. Hyperallometric scaling of reproductive-energy output.
(A) Hyperallometric (b1 = 1.33, solid red curve; see table S7) and isometric
mass scaling (b1 = 1, dashed blue curve) relationships for the Atlantic cod,
G. morhua. Fish sizes span more than one order of magnitude in mass
(2 versus 30 kg). (B) Total population-level biomass (
P
Mi) required to
produce the same reproductive output (
P
Ri, in megajoules) for two
populations of fish with hyperallometric scaling that differ in the mass of the
individual fish, where i is individuals: The standing-biomass ratio differs by
~2.5-fold (that is, 74 versus 30 kg) from the larger individual to the population
with smaller individuals. [Illustration credit: Diane Rome Peebles]
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else being equal, size declines reduce the capacity
of marine fish populations to replenish, particu-
larly if increased egg size increases recruitment
success [but see (15)]. Our findings prove par-
ticularly relevant in light of body-size declines
of major fishery species owing to overharvest-
ing and climate change (10, 16, 17) and suggest
that the reproductive consequences of these size
declines will be dramatic.
REFERENCES AND NOTES
1. R. E. Earll, in National Oceanic and Atmospheric Administration
Report of the United States Commissioner of Fisheries,
S. F. Baird, Ed. (Government Printing Office, San Diego, 1881),
pp. 395–426.
2. R. J. H. Beverton, S. J. Holt, On the Dynamics of Exploited Fish
Populations (Fishery Investigations Series II, Her Majesty’s
Stationery Office, London, 1957), vol. 19.
3. C. T. Marshall, C. L. Needle, A. Thorsen, O. S. Kjesbu,
N. A. Yaragina, Can. J. Fish. Aquat. Sci. 63, 980–994 (2006).
4. M. A. Hixon, D. W. Johnson, S. M. Sogard, ICES J. Mar. Sci. 71,
2171–2185 (2014).
5. J. N. Lim, A. M. Senior, S. Nakagawa, Evolution 68, 2306–2318
(2014).
6. C. Birkeland, P. K. Dayton, Trends Ecol. Evol. 20, 356–358
(2005).
7. S. E. Lester et al., Mar. Ecol. Prog. Ser. 384, 33–46 (2009).
8. Materials and methods are available as supplementary
materials.
9. R. Beldade et al., Proc. R. Soc. Lond. B Biol. Sci. 279, 2116–2121
(2012).
10. I. van Rijn, Y. Buba, J. DeLong, M. Kiflawi, J. Belmaker,
Glob. Chang. Biol. 23, 3667–3674 (2017).
11. D. R. Barneche et al., Ecol. Lett. 17, 1067–1076 (2014).
12. K. H. Andersen et al., ICES J. Mar. Sci. 73, 1651–1655 (2016).
13. R. Froese et al., ICES J. Mar. Sci. 73, 1640–1650 (2016).
14. M. Heino et al., ICES J. Mar. Sci. 70, 707–721 (2013).
15. D. J. Marshall, S. S. Heppell, S. B. Munch, R. R. Warner, Ecology
91, 2862–2873 (2010).
16. C. J. Harvey, N. Tolimieri, P. S. Levin, Ecol. Appl. 16, 1502–1515
(2006).
17. M. J. Genner et al., Glob. Change Biol. 16, 517–527 (2010).
ACKNOWLEDGMENTS
We would like to thank M. Elgar for kindly sharing his life-history
database with us. We thank L. Chapman, B. Comerford, and the
Interlibrary Loan Team at Monash University for helping D.R.B. and
D.J.M. compile the data sets. Many people expended a substantial
amount of effort by providing eggs to D.R.R. for calorimetric
measurements. These included V. Scholey, A. Oliver, E. Schultz,
H. Munehara, P. Sikkel, L. McKelvey, D. Hoesey, and J. Elliott. Numerous
people assisted with laboratory analyses of egg content, including
A. Cedeno, E. Pena, and C. Guerra. People assistingwith field collections
included U. Schober, A. Cedeno, S. Swearer, L. Gutierrez, E. Paredes,
and C. Guerra. We also thank O. J. Luiz for comments on previous
versions of the manuscript. Funding: This study was funded by
the Centre for Geometric Biology, Monash University. Author
contributions: D.R.B., C.R.W., and D.J.M. conceived the study;
D.R.B. and D.J.M. compiled data; D.R.R. collected the egg size and
energy-content data; D.J.M. compiled data on egg dry weight; and
D.R.B. compiled, standardized, and analyzed the data and wrote the
first draft of the manuscript. All authors contributed substantially to
revisions. Competing interests: None declared. Data and materials
availability: All data, metadata, and R code can be downloaded and
cited as “D. R. Barneche, D. R. Robertson, C. R. White, D. J. Marshall,
Data and code from: Fish reproductive-energy output increases
disproportionately with body size. Zenodo (available at https://
github.com/dbarneche/fishFecundity), doi:10.5281/zenodo.1213118.”
SUPPLEMENTARY MATERIALS
www.sciencemag.org/content/360/6389/642/suppl/DC1
Materials and Methods
Fig. S1
Tables S1 to S8
References (18–244)
29 October 2017; accepted 23 March 2018
10.1126/science.aao6868
Barneche et al., Science 360, 642–645 (2018) 11 May 2018 3 of 3
0.5 1.0 1.5 2.0 2.5
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6
11
16
21
26
31
36
41
46
51
56
61
66
71
76
81
86
91
96
101
106
111
Species−specific exponent
0.5 1.0 1.5 2.0 2.5
115
120
125
130
135
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160
165
170
175
180
185
190
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269
274
279
284
289
294
299
304
309
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324
329
334
339
Fig. 3. Species-specific reproductive-energy mass-scaling exponents.
Average exponents and 95% CIs (shown by horizontal bars) for 342
species were obtained by combining species-specific (when available)
posterior estimates from models in Fig. 2, A to C (8). If a species was not
present in a given data set, we used the population-level fixed-effect
posterior estimates for the model in question. The dashed line represents
the reproductive isometry reference point; the dotted line represents the
average fixed-effect exponent value. Red circles and green triangles
represent species whose scaling exponents are greater than and less
than 1, respectively. Fish silhouettes depict the major clades represented
in this figure. See table S7 for species and family names and estimates of
reproductive energy–output slopes.
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Supplementary Materials for 
 
Fish reproductive-energy output increases disproportionately with body size 
 
Diego R. Barneche,* D. Ross Robertson, Craig R. White, Dustin J. Marshall 
 
*Corresponding author. Email: barnechedr@gmail.com 
 
Published 11 May 2018, Science 360, 642 (2018) 
DOI: 10.1126/science.aao6868 
 
 
 
 
This PDF file includes: 
 
Materials and Methods 
Fig. S1 
Tables S1 to S8 
References 
 
 
Materials and Methods
Fecundity and egg size data compilation
We compiled data from multiple previous compilations that provided references on female
fecundity (number of eggs per female) and size, as well as egg size (diameter in mm), for
different species of marine fishes (4, 18–23). We used only raw data from original references
(data extracted from tables or directly from figures using DataThief software). We also
investigated papers cited in these original references, repeating this process exhaustively. We
also searched systematically Google Scholar in English, Spanish and Portuguese using the key
words fish and fecundity, reviewing the first 200 studies returned in each language. We later
separated this compilation into two sub datasets: 1) a dataset containing paired records
between female fecundity and size (n = 7,737 from 134 studies between 1880 to 2015,
including 177 species) and 2) another dataset containing egg size and female size
measurements (n = 926 from 84 studies between 1905 to 2015, including 154 species). A
subset of observations overlapped between these two datasets (i.e. paired measurements of
female size, fecundity, and egg size; n = 1,538, 45 spp.), and were used to analyze the
relationship between total egg-volume (i.e. egg-volume  fecundity, in mm3) and female size
(see Statistical analyses below). We only collected data from studies measuring individuals in
the wild, or individuals that were transferred from the wild to an aquarium facility before the
measurements. For the fecundity dataset, 98.4% of the observations (n = 7,613) were
exclusively from wild-based observations, corresponding to 98.9% (n = 175) of the original
number of species. Removing measurements that were performed in the lab did not alter our
results (Fecundity–Mass model b1 = 1.18; 95% CI: 1.11 – 1.24; see Results section; also see
Statistical analyses below for parameter notation). For the egg size dataset, 96.2% of the
observations (n = 891) were exclusively from wild-based observations, corresponding to
96.8% (n = 149) of the original number of species. Removing measurements that were
performed in the lab did not alter our results (Egg-volume–Mass model b1 = 0.15; 95% CI:
0.07 – 0.23; see Results section). In the egg size dataset, if a study provided multiple body
size measurements for the females but only one population-level average egg size, we took the
median body size across females. For eggs that present ellipsoidal, elongated shapes, we used
both diameters (long, l, and short, s) to first calculate the volume
2
(V = (4=3)p (l=2) (s=2)2), and then back transform to diameter of sphere with
equivalent volume ([((V 3)=(4p))(1=3)]2). All egg size measurements were
transformed to spherical volume, V (mm3; V = (4=3)p r3, where r is radius).
Studies generally reported female length, so we transformed these measured lengths (mm),
l, into mass (g), m, by using the standard length-mass equation m= alb (where a is a
normalization (g cm-b) and b is an exponent that characterizes any changes in shape over
ontogeny). The species-specific length-mass conversion parameters were downloaded from
FishBase (24) (April 2017). For those species with multiple pairs of parameters, we used the
pair with the highest quality score provided in FishBase. For those species that did not have
any parameters, we used an approximation provided by FishBase that is obtained using a
Bayesian method which is based on relationship estimates for the genus-body shape of the
species in question (25).
Quantifying egg energetic content
Fish eggs and data on their volume, dry weight and energy content were collected by DRR
with help from collaborators. A total of 1,111 measurements from 111 species were sampled
following standard techniques (26). Collections took place in Japan, Indonesia, Australia, the
Marquesas Archipelago, Panama, the USA, Canada, and Corsica. Eggs from Panama, USA,
and Canada were collected from both Atlantic and Pacific coasts. Eggs from Corsica were
oven-dried (the lab had no freeze-drier), everything else was freeze dried. Removing the data
from Corsica did not alter our results (Egg-energy–Volume model b1 = 0.77; 95% CI: 0.65 –
0.88; see Results section).
The determination of average dry weight and energy content of an egg in a clutch also
followed standard practice (26). Each sample of eggs was first carefully cleaned of all debris
under a microscope, then rinsed ~6 times in distilled water, and finally stored in a -80˚C
freezer. Subsequently each sample was lyophilized for at least 8 h and stored in a desiccator at
-80˚C. To determine the average dry weight of an egg, a subsample of ~20 counted, intact
dried eggs was weighed on a (daily calibrated) Cahn Model 28 Microbalance, after the
desiccator containing the samples had been allowed to warm to room temperature (~24˚C) in
an air-conditioned room. After this the energy density (Joules mg−1 dry weight) of the sample
3
was determined using a Phillipson Microbomb Calorimeter (Gentry Instruments) calibrated
with a sample of Benzoic acid at the beginning of each day’s set of measurements. Average
energy content of an egg of each clutch was calculated as (average energy density of the
clutch)  (the average dry weight of an egg in that clutch).
We also compiled data from 9 studies that measured both egg-volume and egg-weight (dry
weight in mg; n = 255 observations, 15 spp.). Egg-weight was transformed to energy (Joules)
based on Ref. (27) and our own data (Fig. S1), which show that egg-energy scales
isometrically with egg-weight, with an average of ~25 Joules mg-1 of egg dry weight.
Statistical analyses
Because we are interested in quantifying the mass scaling of fish fecundity and egg size, as
well as the volume scaling of egg-energy content, we applied three linear hierarchical models
with the same structure:
lnY = (lnb0+ lng0spp+ lng0phy)+(b1+ g1spp)lnX+ lne; (1)
where lnY represents a natural log-transformed response vector (fecundity in Fecundity–Mass
model, egg-volume in Egg-volume–Mass model, egg-energy in Egg-energy–Volume model,
and total egg-volume (i.e. fecundity  egg-volume) in Total-volume–Mass), lnb0 is a
fixed-effect intercept, lng0spp and lng0phy are respectively vectors of random-effect coefficients
that account for residual intercept deviations attributable to species uniqueness and patterns of
relatedness as described by the phylogeny, b1 is a fixed-effect slope for the natural
log-transformed predictor vector, lnX (female mass in Fecundity–Mass, Egg-volume–Mass
models and Total-volume–Mass, and egg-volume in Egg-energy–Volume model), g1spp is a
vector of random-effect coefficients that account for residual slope deviations attributable to
species uniqueness, and lne is the model unexplained residual variation.
Phylogenetic relatedness might influence broad-scale variation in life-history traits (28).
From an evolutionary perspective, closely related species might have a higher likelihood of
sharing some ancestral-state trait (29), such as egg-type (pelagic, demersal, brooded). At the
same time, species may present unique variations in traits that are independent of phylogenetic
4
relatedness. From a technical perspective, both of these unmeasured biological factors
(i.e. species uniqueness and relatedness) likely contribute to variance in a particular
life-history trait (30) and, consequently, it is desirable to account for both of these possible
effects. To do so, we created a tree from the Open Tree of Life (OTL) using the rotl R package
(31) v. 3.0.3 in order to test for significant phylogenetic heritability in our models (32). We
first downloaded the full Actinopterygii tree from OTL (n = 38,941 tips) and then added
species from our dataset that were missing in the tree: the Pomacentridae species relationship
followed a recent consensus topology (33) and we inserted them as a sister group to the
Labridae as those are the only two families on the database belonging to Labroidei. One
Centropomidae (Centropomus undecimalis) and one Latidae (Lates calcarifer) were inserted
as a sister group to Carangidae (34). One Apogonidae (Apogon doederleini) was inserted right
next to the other Apogonidae species in the tree. This tree was pruned to retain focal species
only (i.e. individually for each model), and then used to derive a variance-covariance matrix
based on Brownian evolution. The trees included a total of 20, 16, 8, and 2 polytomies for the
Fecundity–Mass, Egg-volume–Mass, Egg-energy–Volume, and Total-volume–Mass models
respectively. Branch lengths are unknown for the phylogeny, so the arbitrary method of
Grafen (35) was applied, whereby branch lengths are set to a length equal to the number of
descendant tips minus one.
In Fecundity–Mass and Total-volume–Mass, fixed effects were assigned informative priors
following a Gaussian distribution (b1: mean = 1, standard deviation = 2; lnb0: mean = 3,
standard deviation = 3) to speed up model convergence. In Egg-volume–Mass and
Egg-energy–Volume, fixed effects were assigned weakly informative priors following a
Gaussian distribution. In all models, random effects were assigned weakly informative priors
following a Gaussian distribution. Random-effect coefficients (lng0spp, lng0phy and g1spp) are
distributed with means of zero, hence representing actual deviations from the fixed effects.
Standard deviations (s

lng0spp

, s

lng0phy

, s

g1spp

and s [lne]) were also assigned weakly
informative priors following a Student-t distribution. We account for the phylogenetic
non-independence among species by weighting the phylogenetic-relatedness standard
deviation (s

lng0phy

) by a variance-covariance matrix, A, which was obtained from the tips
of the phylogeny (i.e. s

lng0phy
A, with A being obtained using the ape R package (36)
5
version 5.0). The phylogenetic heritability (equivalent to Pagel’s l (29, 30)), was estimated as
the proportion of total variance, conditioned on the fixed effects, attributable to the random
effect of phylogeny (i.e. s2

lng0phy

=(s2

lng0phy

+s2

lng0spp

+s2 [lne])). It captured
78.9% (95% CI: 62.2% – 90.2%), 89.3% (95% CI: 80.6% – 95.2%), 98.2% (95% CI: 96.2% –
99.3%), and 5.6% (95% CI: 0.0% – 30.7%) of the variance observed in the residuals in
Fecundity–Mass, Egg-volume–Mass, Egg-energy–Volume, and Total-volume–Mass models
respectively. The posterior distributions of model parameters were estimated using Markov
chain Monte Carlo (MCMC) methods using the brms R package (37) version 2.1.0. For
Fecundity–Mass and Total-volume–Mass, we constructed four chains of 15,000 steps,
including 7,500-step warm-up periods, so a total of 30,000 steps were retained to estimate
posterior distributions (i.e. (15,000 - 7,500)  4 = 30,000). For Egg-volume–Mass and
Egg-energy–Volume, we constructed four chains of 6,000 steps, including 3,000-step warm-up
periods, so a total of 12,000 steps were retained to estimate posterior distributions (i.e. (6,000
- 3,000)  4 = 12,000). Number of chains were tuned in order to achieve model convergence
(i.e. bR= 1).
The dataset used to run Fecundity–Mass includes multiple observations within species,
with clades (e.g. families) having different representation (e.g. many species of the genus
Sebastes, but no other genera within Sebastidae). Thus, it is possible that the average
within-species mass-scaling slope across species would be strongly influenced by a particular
clade. To verify this source of potential bias, we also ran an alternative version of equation 1
whereby we included random slopes attributable to phylogeny-related effects (i.e. adding a
g1phy to the model). This alternative model was compared to the simpler Fecundity–Mass
model using leave-one-out cross-validation (LOO), which, similarly to widely applicable
information criterion (WAIC), is a fully Bayesian model selection procedure for estimating
pointwise out-of-sample prediction accuracy (38, 39). For each model, we calculated the
expected log pointwise predictive density (delpdloo) using the log-likelihood evaluated at the
posterior simulations of the parameter values (39). We calculated p-values for the pairwise
differences in delpdloo (Ddelpdloo) using standard errors (s.e.) and a normal probability density
function. This method of calculating s.e.’s is reliable for data sets with many observations (n =
7,737 in our analysis) because the distribution of Ddelpdloo is well approximated by a normal
6
distribution (39). Although the alternative model had higher average predictive accuracy than
Fecundity–Mass, it was not significantly higher (p = 0.4498), and we therefore retained
Fecundity–Mass, which is simpler. We also compared Fecundity–Mass, Egg-volume–Mass,
and Egg-energy–Volume models each against a more complex version which included an
interaction between scaling slope and spawning mode (categorical fixed effect with 6 levels
added to Fecundity–Mass: demersal, internal brooding, mouth brooder, pelagic, pouch
brooder, scatterer; 6 levels added to Egg-volume–Mass: demersal, internal brooding, mouth
brooder, pelagic, pouch brooder, scatterer; 3 levels added to Egg-energy–Volume: demersal,
mouth brooder, pelagic) to test for an effect of spawning mode on the estimated scaling slopes
after accounting for phylogenetic effects. However, in our three datasets, spawning mode and
phylogeny are strongly confounded, and adding an interaction term did not improve model fit
significantly for any of the three models (p = 0.9116 in Fecundity–Mass; p = 0.5492 in
Egg-volume–Mass; p = 0.3053 in Egg-energy–Volume). Model comparison was implemented
using the loo R package (39) version 1.1.0.
Estimating total reproductive-energy output
We estimated the average mass scaling of total reproductive-energy output per spawning per
female by combining the estimated responses from our three Bayesian models. Specifically,
considering that Fecundity–Mass, Egg-volume–Mass, and Egg-energy–Volume models can be
translated into
Fecundity= b0M1Massb1M1 (2)
Egg-volume= b0M2Massb1M2 (3)
Egg-energy= b0M3Egg-volumeb1M3 ; (4)
where b0 and b1 respectively represent average fixed-effect intercepts and slopes from Models
M13, one can first predict the expected egg-energy for a female of given mass,
7
Egg-energy= b0M3bb1M30M2 Mass
b1M2b1M3 (5)
and then multiply this expectation by the expected fecundity for the same given mass, yielding
Total reproductive-energy output= b0M1b0M3bb1M30M2 Mass
b1M1+(b1M2b1M3) (6)
This estimate was calculated 10,000 times by randomly sampling from each of the models’
posterior distributions in order to fully propagate each parameter uncertainty. We note that this
approach is very conservative, as the parameter estimates from each model are completely
independent from each other (i.e. it is not possible to compute the covariance among
parameters from different models) and therefore the total propagated error is likely to be
overinflated. Thus, if the compound 95% credible intervals for the mass scaling slope are
indeed greater than 1 we can safely assume that the average total reproductive-energy output
scales super-linearly with body mass.
Exploring subsets of the data for which we have overlapping metrics
We used statistical methods to generate total reproductive-energy output because, for most
species, we did not have all four metrics of interest (i.e. body size, fecundity, egg size, and egg
energy). However, for 45 species (n = 1,538), we had both female size-fecundity, and female
size-egg size data and so we could estimate the female size-total reproductive volume
relationship for that subset directly. Like the complete dataset, the analysis of this subset
revealed hyper-allometric scaling between female size and total reproductive volume where
the predicted exponent was greater than 1 (1.21; 95% CI: 1.07 – 1.37; Table S6). For 9
species, we had data on all three relationships of interest and for that restricted group, the
mean scaling exponent between female size and reproductive energy output was again
hyper-allometric: 1.18 (see species marked with an asterisk in Table S7). Finally, for just 3
species, we had data on all four variables of interest (female size, fecundity, egg-volume, and
egg-energy) where all four came from the same study and population. For this very small
group, we again found evidence of hyper-allometric scaling in two of the three species and
isometry for the third species (Table S8).
8
Calculating reproductive loss: MPAs and ocean warming
In the main text, we show that, for the widow rockfish Sebastes entomelas, the benefits of an
MPA to population replenishment for fecundity and reproductive-energy output are
respectively 60% and 74% under hyper-allometric scaling. These numbers were calculated
assuming a fixed standing biomass of 1 tonne both inside and outside an MPA, a baseline size
of 30 cm outside the MPA (~350 g), and a size of 38.4 cm (i.e. 28% larger in length (7)) inside
the MPA (~748 g). Thus, we have 2,857 individuals outside vs. 1,337 individuals inside the
MPA. The hyper-allometric species-specific fecundity (b0 = 0.19, b1 = 1.62; Table S3) and
reproductive-energy output (b0 = 220.14, b1 = 1.73; Table S7) parameters for this species
were used to calculate both the total fecundity and reproductive-energy output outside
(fecundity = 2,857  0.19  3501.62 = 7,178,837 eggs; reproductive-energy = 2,857  220.14
 3501.73 = 15,843,346,454 J) and inside (fecundity = 1,337  0.19  7481.62 = 11,497,558
eggs; reproductive-energy = 1,337  220.14  7481.73 = 27,585,453,858 J) the MPA.
Therefore, (11,497,558 / 7,178,837 - 1)  100  60%, and (27,585,453,858 / 15,843,346,454
- 1)  100  74%.
We also showed that a 15% decrease in fish length due to a warming of 1.5˚C (10) would
incur a 50% per-capita reduction in fecundity for Atlantic mackerel, Scomber scombrus.
These were calculated assuming a baseline size of 50 cm before (~925 g), and a size of 42.5
cm (i.e. 15% smaller, ~561 g) after warming. The species-specific fecundity ( b0 = 3.73, b1 =
1.38; Table S3) parameters for this species were used to calculate fecundity before (3.73 
9251.38 = 46,236 eggs) and after (3.73  5611.38 = 23,189 eggs) warming. Therefore, (1 -
23,189 / 46,236)  100  50%.
Relationship between egg-energy and female size
In the main text, we state that the combination of the relationship between egg-energy and
egg-volume with the relationship between egg-volume and female mass yields a positive,
hypo-allometric relationship between female size and per-egg energy content. This is because
egg-energy µ mass1.18  0.77  mass 0.91 (equation 5).
9
Fig. S1. Relationship between egg energy (J) and egg dry weight (mg). All data collected
experimentally by D. Ross Robertson.
10
Table S1. Summary of models that make explicit predictions or assumptions regarding
the relationship between female body size and reproductive output. Reproductive output is
characterized in a variety of ways in these models, from fraction of the growing season de-
voted to reproduction to ratios of gonad mass to somatic mass. * indicates studies that make
explicit assumptions about the energy content of reproductive materials and how that scales
with body size (all assume isometric scaling of energy content). FGO and VGO = Fixed and
Variable (respectively) allocation to reproduction.
Study Model type Scaling Prediction or Assumption Refs
Gadgil and Bossert (1970) Life history Hyperallometric Prediction (40)
Roff (1983) Life history Isometric Assumption (41)
Roff (1984) Life history Isometric Assumption (42)
Reiss (1985) Life history Hypoallometric Prediction (43)
Kozłowski and Uchmanksi (1987) Life history Variable Prediction (44)
Kozłowski (1996) Life history Variable Prediction (45)
West et al. (2001) Life history Isometric Assumption (46)
Charnov et al. (2001) Life history Isometric Assumption (47)
Charnov (2002) Life history Isometric Assumption (48)
Lester et al. (2004) Life history Isometric* Assumption (49)
Roff et al. (2006) Life history Isometric Assumption (50)
Quince et al. (2008; FGO) Life history Isometric* Assumption (51)
Quince et al. (2008; VGO) Life history Hyperallometric* Prediction (51)
Pecquerie et al. (2009) Life history Isometric Assumption (52)
Koojiman (2010) Life history Isometric* Assumption (53)
Arendt (2011) Life history Isometry Assumption (54)
Onishi et al. (2012) Life history Hyperallometric Assumption (55)
Brunel et al. (2013) Life history Isometric Assumption (56)
Charnov et al. (2013) Life history Isometric Assumption (57)
Boukal et al. (2014) Life history Isometric* Assumption (58)
Koojiman and Lika (2014) Life history Isometric* Assumption (59)
Minte-Vera et al. (2016) Life history Isometric* Assumption (60)
Jusup et al. (2017) Life history Isometric Assumption (61)
Mangel (2017) Life history Hyperallometric Assumption (62)
Smallegange et al. (2017) Life history Isometric Assumption (63)
Beverton and Holt (1957) Fisheries Isometric Assumption (2)
Scott et al. (2006) Fisheries Hyperallometric Assumption (64)
Jørgensen and Fisken (2006) Fisheries Isometric* Assumption (65)
Enberg et al. (2010) Fisheries Isometric Assumption (66)
Eikeset et al. (2013) Fisheries Isometric Assumption (67)
Lester et al. (2014) Fisheries Isometric Assumption (68)
Andersen and Beyer (2015) Fisheries Isometric Assumption (69)
Eikeset et al. (2016) Fisheries Isometric Assumption (70)
Andersen et al. (2016) Fisheries Isometric Assumption (12)
Zimmerman and Jørgensen (2016) Fisheries Isometric Assumption (71)
Hartvig et al. (2011) Food web Isometric Assumption (72)
Carozza et al. (2016) Food web Isometric Assumption (73)
11
Table S2. Summary table of the major compilations of fish reproductive scaling relation-
ships (both within and among species). Note that where scaling relationships used length
rather than mass, we assumed that mass scaled to length3.
Study Interspecific/intraspecific Relationship Allometric Scaling Refs
Blueweiss et al. (1978) Inter- Size-fecundity Hypo- (74)
Blueweiss et al. (1978) Inter- Size-egg size Hypo- (74)
Wootton (1979) Intra- Size-fecundity Hyper- (75)
Roff (1983) Intra- Size-fecundity Hyper- (41)
Duarte and Alcaraz (1989) Inter- Size-fecundity Hypo- (19)
Duarte and Alcaraz (1989) Inter- Size-egg size None (19)
Wootton (1992) Inter- Size-egg size Iso- (76)
Hayward and Gillooly (2011) Inter- Size-fecundity Iso- (77)
Hixon et al. (2014) Intra- Size-fecundity Hyper- (4)
Dick et al. (2017) Intra- Size-fecundity Hyper- (78)
12
Table S3. Summary of data used for Fecundity–Mass model. This table shows summary stats of raw data used in Fecundity–Mass model (Fig. 2A). OTL =
Open Tree of Life Identifier; SM = Spawning Mode (D = demersal, P = pelagic, S = scatterer, BM/BP/BI = mouth/pouch/internal brooder – all brooders have no
parental provisioning post fertilization); F3 = Number ID in Fig. 3; b1 / 2.5% / 97.5% = Estimates (mean and 95% Bayesian credible intervals) of species-specific
fecundity mass-scaling exponents as obtained from Fecundity–Mass model (see Materials and Methods); n = number of independent observations for each species.
Family Species OTL SM F3 b1 2.5% C.I. 97.5% C.I. n Fecundity range Mass range (g) Refs
Agonidae Leptagonus decagonus ott3632024 D 269 1.16 0.66 1.67 1 577 32.32 (79)
Apogonidae Cheilodipterus macrodon ott965176 BM 59 1.27 0.71 1.83 1 2,600 110.09 (80)
Apogonidae Holapogon maximus ott3635086 BM 61 1.24 0.71 1.77 1 1,800 48.48 (80)
Apogonidae Jaydia hungi ott3634955 BM 62 1.23 0.73 1.73 1 800 20.36 (80)
Apogonidae Ostorhinchus cookii ott102070 BM 63 1.25 0.76 1.74 1 500 15.91 (80)
Apogonidae Ostorhinchus cyanosoma ott638185 BM 64 1.09 0.65 1.53 1 650 6.02 (80)
Apogonidae Siphamia tubifer ott5860863 BM 67 0.86 0.52 1.19 1 240 0.29 (80)
Apogonidae Taeniamia lineolata ott775407 BM 68 1.07 0.65 1.49 1 400 3.35 (80)
Atherinidae Atherina presbyter ott669918 D 2 1.09 0.9 1.28 33 935 – 11,407 1.47 – 13.70 (81)
Atherinopsidae Leuresthes tenuis ott274466 D 3 1.39 0.93 1.87 7 475 – 2,705 15.99 – 31.31 (82)
Atherinopsidae Odontesthes argentinensis ott93287 D 6 1.17 0.86 1.48 24 3,422 – 27,980 41.62 – 244.65 (83)
Balistidae Balistes capriscus ott135140 S 335 1.61 1.42 1.8 172 32,919 – 1,996,500 192.48 – 1002.07 (84); (85)
Blenniidae Aidablennius sphynx ott898527 D 70 1.07 0.68 1.46 1 655 0.63 (86)
Bothidae Arnoglossus laterna ott696263 P 258 0.98 0.6 1.37 2 7,092 – 59,574 2.68 – 9.09 (87)
Carangidae Decapterus punctatus ott705034 P 74 1.28 0.84 1.72 2 6,200 – 51,000 16.71 – 53.59 (88)
Carangidae Elagatis bipinnulata ott705026 P 75 0.99 0.73 1.25 29 469,590 – 2,635,300 1378.72 – 7075.53 (89)
Carangidae Trachurus picturatus ott174574 P 77 1.13 0.65 1.62 8 23,630 – 48,816 38.10 – 73.11 (90)
Centropomidae Centropomus undecimalis ott317368 P 79 1.49 1.08 1.9 15 578,160 – 10,289,069 2711.08 – 7993.41 (91)
Clupeidae Amblygaster sirm ott328216 P 11 1.57 1.13 2.04 33 20,281 – 140,411 34.39 – 57.50 (92)
Clupeidae Clupea harengus ott1005932 D 14 1.54 1.42 1.66 419 19,355 – 265,161 140.27 – 533.37 (93)
Clupeidae Ethmalosa fimbriata ott658559 P 16 0.89 0.56 1.23 32 15,816 – 51,901 182.55 – 582.47 (94)
Clupeidae Nematalosa vlaminghi ott328199 P 17 1.15 0.59 1.71 1 90,451 111.79 (95)
Clupeidae Opisthonema libertate ott437031 P 18 1.1 0.7 1.5 58 22,025 – 106,958 91.39 – 204.30 (96)
Clupeidae Opisthonema medirastre ott1195 P 19 1.24 0.82 1.67 46 17,941 – 107,063 80.90 – 169.98 (97)
Clupeidae Sardinops sagax ott121238 P 20 0.97 0.83 1.11 185 3,083 – 33,483 26.53 – 118.21 (98)
Clupeidae Spratelloides gracilis ott39732 D 21 0.99 0.74 1.26 18 594 – 5,913 0.20 – 0.86 (99)
Clupeidae Strangomera bentincki ott648871 P 22 1.08 0.65 1.49 10 5,901 – 10,414 11.25 – 24.31 (100)
Cottidae Artediellus atlanticus ott654509 D 273 1.32 0.82 1.81 1 88.5 23.32 (79)
Cottidae Enophrys bison ott290389 D 274 1.13 0.55 1.7 2 18,800 – 31,900 197.53 – 241.50 (101)
Cottidae Icelus bicornis ott3632116 D 276 1.1 0.6 1.59 1 440 14.79 (79)
Cottidae Triglops pingelii ott568671 D 277 1.12 0.63 1.62 1 600 22.25 (79)
Cyclopteridae Cyclopteropsis mcalpini ott3631993 D 279 1.04 0.63 1.46 1 65 2.80 (79)
Engraulidae Anchoviella lepidentostole ott610213 P 23 0.97 0.59 1.35 24 14,395 – 37,191 9.36 – 21.80 (102)
Engraulidae Cetengraulis mysticetus ott1064953 P 24 1.23 0.98 1.47 86 9,810 – 79,010 25.10 – 71.50 (103)
Engraulidae Engraulis anchoita ott648864 P 25 1.26 1.12 1.4 159 1,641 – 36,464 5.44 – 39.01 (104)
Engraulidae Engraulis mordax ott29754 P 26 1.16 0.76 1.56 21 8,673 – 35,561 10.47 – 23.98 (105)
Engraulidae Engraulis ringens ott211981 P 27 1.35 1.14 1.56 149 3,067 – 34,977 10.78 – 39.92 (106); (100); (107)
Exocoetidae Hirundichthys affinis ott837204 D 7 1.21 0.63 1.79 2 7,398 – 10,021 218.16 – 255.62 (108)
Gadidae Boreogadus saida ott877055 P 33 0.89 0.41 1.38 1 48,500 18.48 (79)
Gadidae Eleginus nawaga ott450082 D 34 1.12 0.57 1.66 1 40,500 74.86 (79)
Gadidae Gadus morhua ott5860467 P 35 1.23 1.18 1.28 428 151,700 – 21,496,000 1036.67 – 24086.31 (109); (110); (111); (112)
Gadidae Merlangius merlangus ott635911 P 37 1.03 0.92 1.15 178 95,555 – 1,443,958 94.88 – 1467.11 (113)
13
Table S3 Continued.
Family Species OTL SM F3 b1 2.5% C.I. 97.5% C.I. n Fecundity range Mass range (g) Refs
Gadidae Micromesistius australis ott1091056 P 38 1.56 1.37 1.76 96 2,658 – 81,300 188.76 – 1017.03 (114)
Gasterosteidae Gasterosteus aculeatus ott111122 D 45 0.91 0.78 1.04 110 48 – 523 0.33 – 3.03 (115); (116)
Gobiidae Elacatinus oceanops ott638369 D 89 0.8 0.44 1.15 1 375 0.25 (117)
Gobiidae Gobiosoma robustum ott1064480 D 90 1.35 0.76 1.95 1 695 158.08 (118)
Gobiidae Paragobiodon echinocephalus ott940827 D 92 0.87 0.5 1.24 1 716 0.23 (119)
Gobiidae Paragobiodon lacunicolus ott758973 D 93 0.88 0.53 1.24 1 635 0.23 (119)
Gobiidae Paragobiodon xanthosoma ott770378 D 94 0.96 0.61 1.32 1 293 0.23 (119)
Gobiidae Pomatoschistus minutus ott1048873 D 95 0.82 0.46 1.17 28 1,430 – 4,070 1.53 – 3.11 (120)
Haemulidae Haemulopsis corvinaeformis ott177515 P 97 0.52 0.3 0.75 2 15,056 – 83,316 0.93 – 51.66 (108)
Hemiramphidae Hemiramphus brasiliensis ott953384 D 8 0.84 0.44 1.24 2 2,200 – 4,000 21.01 – 111.47 (108)
Hemitripteridae Hemitripterus americanus ott160287 D 282 1.22 0.78 1.66 6 1,039 – 22,194 303.44 – 1004.16 (121)
Hexagrammidae Oxylebius pictus ott131554 D 284 0.92 0.44 1.41 22 12,789 – 25,621 32.22 – 63.30 (122)
Labridae Thalassoma bifasciatum ott907850 P 145 0.75 0.6 0.9 66 79 – 5,140 0.52 – 4.60 *
Latidae Lates calcarifer ott6362446 P 150 1.04 0.83 1.26 25 2,062,100 – 47,202,800 2945.25 – 32293.52 (123)
Liparidae Careproctus reinhardti ott1065176 D 285 1.23 0.62 1.83 1 300 357.25 (79)
Liparidae Paraliparis bathybius ott309222 BM 287 1.25 0.68 1.82 1 428 122.99 (79)
Lutjanidae Lutjanus campechanus ott642863 P 151 1 0.62 1.37 9 160,000 – 900,000 1350.00 – 5850.00 (124)
Lutjanidae Lutjanus carponotatus ott333348 P 152 1.72 1.51 1.93 55 4,451 – 750,512 77.08 – 485.66 (125)
Lutjanidae Lutjanus synagris ott943184 P 153 0.67 0.46 0.87 39 23,000 – 160,000 198.00 – 2100.00 (126)
Lutjanidae Ocyurus chrysurus ott943183 P 154 1.25 0.88 1.61 21 14,268 – 166,651 221.70 – 734.70 (127)
Lutjanidae Rhomboplites aurorubens ott642869 P 155 1.39 1.24 1.53 41 4,350 – 1,792,070 158.61 – 2994.94 (128)
Malacanthidae Caulolatilus microps ott39883 P 156 1.47 1.22 1.73 43 83,470 – 4,120,570 762.87 – 4527.12 (129)
Merlucciidae Merluccius gayi ott323082 P 42 1.19 0.9 1.48 50 45,041 – 399,890 523.49 – 3390.80 (130)
Merlucciidae Merluccius hubbsi ott757316 P 43 1.06 0.99 1.14 251 37,817 – 2,537,392 261.48 – 5782.35 (131)
Merlucciidae Merluccius merluccius ott5860466 P 44 0.93 0.77 1.1 209 32,665 – 578,510 626.18 – 3465.51 (132)
Monacanthidae Brachaluteres jacksonianus ott333693 S 336 1.2 0.83 1.56 1 114 1.24 (133)
Monacanthidae Eubalichthys bucephalus ott3631310 S 337 1.32 0.73 1.91 2 9,000 – 39,500 355.86 – 524.85 (134)
Monacanthidae Oxymonacanthus longirostris ott333706 S 338 1.31 0.91 1.72 3 100 – 200 3.30 – 3.79 (135)
Monacanthidae Rudarius ercodes ott511593 S 340 0.83 0.48 1.17 3 2,560 – 8,180 1.25 – 5.78 (136)
Moronidae Dicentrarchus labrax ott3549 P 157 1.23 0.85 1.6 16 290,390 – 2,043,126 1000.00 – 3800.00 (137)
Mugilidae Chelon labrosus ott777079 P 46 1.09 0.58 1.61 13 353,000 – 745,000 1240.00 – 2300.00 (138)
Mugilidae Chelon ramada ott668241 P 47 1.18 0.62 1.73 2 581,000 – 1,243,000 850.00 – 1450.00 (138)
Mugilidae Mugil cephalus ott418648 P 48 1.01 0.93 1.1 95 43,182 – 4,800,000 42.77 – 3760.17 (139); (140); (141)
Mugilidae Mugil curema ott418643 P 49 1.15 0.6 1.69 8 295,081 – 488,956 222.87 – 371.93 (142)
Mugilidae Mugil hospes ott779937 P 50 1.14 0.58 1.7 1 178,277 114.11 (142)
Mugilidae Mugil liza ott779933 P 51 1.19 0.61 1.75 1 1,776,309 1201.53 (142)
Mugilidae Mugil trichodon ott246901 P 52 1.18 0.62 1.73 1 100,346 112.33 (142)
Mugilidae Planiliza subviridis ott823526 P 53 1.07 0.66 1.48 12 38,167 – 145,883 11.69 – 28.41 (143)
Nototheniidae Lepidonotothen nudifrons ott260052 D 160 1.27 0.69 1.84 1 2,500 117.20 (144)
Osmeridae Mallotus villosus ott225330 D 54 1.22 0.7 1.74 34 6,700 – 14,300 18.07 – 26.43 (145)
Paralichthyidae Paralichthys dentatus ott1019735 P 259 1.06 0.91 1.2 134 460,059 – 4,202,538 2627.81 – 20166.83 (146)
Paralichthyidae Paralichthys patagonicus ott654479 P 260 1.38 0.91 1.85 24 20,084 – 156,527 709.32 – 1465.65 (147)
Plesiopidae Acanthoclinus fuscus ott3633802 D 164 1.46 0.94 2.01 7 42,756 – 438,961 232.38 – 475.62 (148)
Pleuronectidae Hippoglossoides platessoides ott223665 P 262 0.96 0.83 1.09 154 25,700 – 258,000 24.83 – 255.51 (149); (150)
Pleuronectidae Pleuronectes platessa ott408337 P 264 1.11 1.04 1.18 486 28,800 – 754,600 189.13 – 3022.59 (151)
14
Table S3 Continued.
Family Species OTL SM F3 b1 2.5% C.I. 97.5% C.I. n Fecundity range Mass range (g) Refs
Pleuronectidae Pseudopleuronectes americanus ott261419 D 265 0.99 0.63 1.35 17 192,240 – 1,390,495 332.81 – 1322.71 (152)
Pleuronectidae Reinhardtius hippoglossoides ott835141 P 266 1.3 1.07 1.53 88 6,761 – 70,394 970.70 – 5331.71 (153)
Pomacanthidae Pomacanthus zonipectus ott826472 P 165 1.25 0.69 1.81 5 25,456 – 62,401 160.12 – 211.09 (154)
Pomacentridae Abudefduf saxatilis ott405751 D 167 0.94 0.43 1.44 1 31,088 22.12 (155)
Pomacentridae Acanthochromis polyacanthus ott100410 D 172 1.41 0.85 1.96 15 245.6 – 573.1 87.19 – 124.17 (156)
Pomacentridae Pomacentrus coelestis ott622060 D 180 1.02 0.75 1.29 81 1,391 – 6,213 1.27 – 3.26 (157)
Pomacentridae Stegastes fuscus ott3635543 D 185 0.96 0.63 1.3 14 1,759 – 14,856 3.92 – 14.19 (158)
Pristigasteridae Ilisha africana ott741177 P 28 1.3 0.78 1.82 1 5,227 37.69 (159)
Sciaenidae Bairdiella chrysoura ott761907 P 207 1.09 0.56 1.62 1 52,800 54.91 (160)
Sciaenidae Cynoscion regalis ott5860595 P 208 1.49 1.24 1.73 29 90,019 – 2,163,872 554.68 – 3969.46 (161)
Sciaenidae Cynoscion striatus ott267367 P 209 1.23 0.66 1.79 5 45,347 – 98,573 178.46 – 230.71 (162)
Sciaenidae Genyonemus lineatus ott1027240 P 210 1.9 1.66 2.15 44 1,038 – 36,701 78.67 – 401.10 (163)
Sciaenidae Isopisthus parvipinnis ott133138 P 211 1.31 0.88 1.76 8 4,280 – 51,604 19.18 – 44.36 (162)
Sciaenidae Larimus breviceps ott924070 P 212 1.71 1.2 2.25 6 5,463 – 165,132 73.67 – 144.63 (162)
Sciaenidae Larimus fasciatus ott897432 P 213 1.03 0.86 1.2 81 17,639 – 315,972 8.75 – 59.86 (164)
Sciaenidae Macrodon ancylodon ott99076 P 215 1.24 0.89 1.59 21 26,210 – 217,632 155.00 – 500.00 (165)
Sciaenidae Micropogonias furnieri ott742890 P 216 0.99 0.83 1.16 85 44,733 – 776,006 315.49 – 2485.35 (166); (167)
Sciaenidae Paralonchurus brasiliensis ott160812 P 218 0.89 0.42 1.35 16 11,497 – 86,034 44.15 – 94.52 (162)
Sciaenidae Seriphus politus ott827210 P 219 1.11 1 1.22 143 2,314 – 125,632 7.44 – 85.10 (168)
Sciaenidae Stellifer rastrifer ott362452 P 220 0.76 0.5 1.02 44 1,544 – 263,392 21.73 – 107.90 (162)
Scombridae Auxis rochei ott713972 P 221 1.18 0.59 1.78 1 75,000 453.17 (169)
Scombridae Scomber scombrus ott70561 P 225 1.38 1.08 1.68 27 151,916 – 1,491,192 190.50 – 832.39 (170)
Scombridae Scomberomorus cavalla ott70546 P 226 1.37 1.26 1.49 75 69,264 – 12,206,888 886.37 – 28991.83 (171); (172)
Scombridae Scomberomorus maculatus ott266387 P 227 1.12 0.84 1.4 14 525,000 – 7,411,000 550.00 – 4150.00 (173); (1)
Scombridae Thunnus alalunga ott833194 P 228 0.93 0.56 1.32 69 263,100 – 2,813,400 14094.95 – 25217.33 (174)
Scombridae Thunnus albacares ott833188 P 229 0.99 0.57 1.4 4 1,300,000 – 5,431,000 32593.36 – 57119.99 (175)
Scombridae Thunnus thynnus ott358607 P 231 1.31 0.83 1.79 27 13,600,000 – 57,600,000 1336.98 – 2861.01 (176)
Sebastidae Sebastes alutus ott480735 BI 288 1.76 1.6 1.93 88 4,500 – 270,000 255.24 – 1429.15 (177); (178)
Sebastidae Sebastes atrovirens ott480732 BI 289 1.11 0.59 1.62 17 67,927 – 528,987 268.54 – 471.51 (148)
Sebastidae Sebastes auriculatus ott480729 BI 290 1.06 0.74 1.39 35 51,619 – 338,540 391.43 – 1455.21 (179)
Sebastidae Sebastes brevispinis ott1057081 BI 291 1.28 0.81 1.76 22 496,000 – 1,252,000 1448.00 – 2832.00 (180)
Sebastidae Sebastes carnatus ott1057074 BI 292 1.19 0.67 1.7 11 77,671 – 236,038 499.24 – 1054.02 (148)
Sebastidae Sebastes caurinus ott1057078 BI 293 1.68 1.49 1.86 54 15,642 – 640,052 247.45 – 2343.44 (179); (181)
Sebastidae Sebastes chlorostictus ott1057068 BI 294 1.55 1.34 1.76 64 17,198 – 761,362 246.06 – 1252.35 (182); (183)
Sebastidae Sebastes constellatus ott1057062 BI 295 1.34 0.95 1.73 21 31,781 – 248,490 224.89 – 655.33 (182)
Sebastidae Sebastes crameri ott1057065 BI 296 1.46 1.1 1.82 12 36,600 – 609,800 491.77 – 2582.59 (184)
Sebastidae Sebastes dallii ott1088425 BI 297 1.15 0.75 1.55 23 3,744 – 18,131 17.90 – 44.78 (182)
Sebastidae Sebastes diploproa ott1057059 BI 298 1.45 1.13 1.77 15 14,400 – 303,700 85.27 – 559.21 (184)
Sebastidae Sebastes elongatus ott531065 BI 299 1.3 1.06 1.53 25 8,707 – 296,799 56.35 – 567.53 (182)
Sebastidae Sebastes entomelas ott531068 BI 300 1.62 1.43 1.81 110 55,600 – 1,102,844 443.86 – 2398.20 (185); (182); (184)
Sebastidae Sebastes flavidus ott531059 BI 301 1.5 1.25 1.74 49 48,400 – 969,715 395.33 – 2310.74 (184); (182)
Sebastidae Sebastes goodei ott531047 BI 302 1.32 1.13 1.5 62 29,200 – 961,809 351.87 – 2426.91 (182); (184)
Sebastidae Sebastes helvomaculatus ott531044 BI 303 1.29 0.81 1.79 5 183,000 – 1,652,000 105.65 – 322.55 (186)
Sebastidae Sebastes hopkinsi ott9589 BI 304 1.01 0.66 1.36 39 8,332 – 39,559 61.99 – 228.24 (182)
Sebastidae Sebastes jordani ott9592 BI 305 1.1 0.75 1.47 10 6,200 – 50,100 28.36 – 168.04 (184)
15
Table S3 Continued.
Family Species OTL SM F3 b1 2.5% C.I. 97.5% C.I. n Fecundity range Mass range (g) Refs
Sebastidae Sebastes levis ott9598 BI 306 1.06 0.79 1.34 27 181,001 – 1,941,579 1367.68 – 7660.85 (182)
Sebastidae Sebastes melanops ott9604 BI 307 1.3 1.05 1.54 226 214,367 – 1,498,862 1165.81 – 2799.49 (187)
Sebastidae Sebastes melanostomus ott9607 BI 308 1.41 1.11 1.73 42 174,874 – 1,177,889 864.30 – 2919.60 (188); (186)
Sebastidae Sebastes mentella ott1088432 BI 309 1.46 1.37 1.55 227 1,457 – 115,846 162.42 – 1284.55 (189); (190)
Sebastidae Sebastes miniatus ott9611 BI 310 1.58 1.37 1.78 59 63,300 – 2,647,282 1022.07 – 10240.11 (182); (184)
Sebastidae Sebastes mystinus ott9616 BI 311 1.74 1.58 1.9 151 10,369 – 524,554 192.42 – 983.32 (148); (191)
Sebastidae Sebastes norvegicus ott1088441 BI 312 1.06 0.89 1.23 67 19,810 – 336,700 434.15 – 3204.64 (192); (190)
Sebastidae Sebastes ovalis ott582324 BI 313 1.23 0.82 1.64 39 52,356 – 212,399 425.50 – 1019.60 (186); (188)
Sebastidae Sebastes paucispinis ott582321 BI 314 1.35 1.18 1.53 75 20,000 – 2,297,900 724.86 – 6193.24 (182); (184)
Sebastidae Sebastes pinniger ott582318 BI 315 1.26 0.79 1.73 10 260,600 – 1,897,600 1506.08 – 4055.36 (184)
Sebastidae Sebastes rastrelliger ott582315 BI 316 1.23 0.82 1.65 2 80,000 – 760,000 386.67 – 2211.98 (193)
Sebastidae Sebastes rosaceus ott212767 BI 317 1.14 0.84 1.44 23 12,992 – 94,936 43.53 – 172.48 (182)
Sebastidae Sebastes rosenblatti ott582311 BI 318 1.29 0.97 1.61 26 4,880 – 93,370 47.04 – 171.78 (182)
Sebastidae Sebastes rufus ott884513 BI 319 1.3 0.93 1.67 27 67,051 – 606,888 672.05 – 2143.58 (182)
Sebastidae Sebastes saxicola ott884510 BI 320 1.04 0.94 1.14 43 4,376 – 230,000 13.98 – 452.33 (182); (184)
Sebastidae Sebastes semicinctus ott795707 BI 321 1.29 0.99 1.61 46 3,275 – 30,867 18.95 – 85.87 (182)
Sebastidae Sebastes serranoides ott884519 BI 322 1.56 1.34 1.78 83 29,597 – 524,048 287.54 – 1198.45 (194)
Sebastidae Sebastes viviparus ott99016 BI 323 1.21 0.94 1.49 32 1,770 – 29,047 69.51 – 324.99 (195)
Serranidae Cephalopholis cruentata ott12880 P 232 1.18 0.7 1.66 12 86,204 – 615,387 142.95 – 414.81 (196)
Serranidae Epinephelus aeneus ott306171 P 233 1.15 0.87 1.42 27 404,056 – 12,589,242 967.53 – 6975.90 (197)
Serranidae Epinephelus fasciatus ott534146 P 234 0.78 0.34 1.21 9 254,732 – 879,038 395.33 – 1435.81 (197)
Serranidae Epinephelus marginatus ott733897 P 235 1.22 0.65 1.78 2 257,657 – 606,246 758.70 – 1197.83 (197)
Serranidae Epinephelus tauvina ott218649 P 236 0.9 0.51 1.29 4 850,186 – 2,904,912 1062.45 – 6012.89 (198)
Serranidae Mycteroperca tigris ott207413 P 237 1.43 0.95 1.93 2 890,000 – 10,490,000 3130.04 – 7720.75 (199)
Siganidae Siganus canaliculatus ott199059 S 241 1.25 1.05 1.44 27 35,494 – 1,555,688 59.67 – 870.20 (200)
Soleidae Solea solea ott710233 P 268 1.15 1.07 1.24 323 50,400 – 1,577,100 117.75 – 1194.75 (201)
Sparidae Acanthopagrus latus ott760733 P 242 1.16 0.58 1.73 5 1,362,137 – 2,152,993 426.21 – 660.47 (202)
Sparidae Archosargus rhomboidalis ott724892 P 243 0.91 0.69 1.13 23 18,274 – 161,107 15.05 – 156.17 (203)
Sparidae Sparidentex hasta ott845300 P 244 1.21 0.73 1.69 6 308,273 – 1,693,365 2604.06 – 6288.57 (202)
Stromateidae Pampus chinensis ott916290 P 245 1.14 0.56 1.71 8 80,400 – 93,100 647.18 – 876.77 (204)
Syngnathidae Hippocampus reidi ott630138 BP 326 1.24 0.76 1.72 3 725 – 914 10.00 – 12.30 (205)
Syngnathidae Micrognathus crinitus ott3642321 BP 327 1.11 0.79 1.43 1 21.7 0.17 (206)
Syngnathidae Syngnathus floridae ott774435 BP 330 1.16 0.74 1.58 1 357.6 3.03 (206)
Syngnathidae Syngnathus louisianae ott774414 BP 331 1.22 0.77 1.69 1 664.7 8.36 (206)
Syngnathidae Syngnathus scovelli ott251400 BP 333 1.2 0.83 1.58 1 55.6 0.88 (206)
Tetraodontidae Canthigaster valentini ott701127 S 342 0.97 0.77 1.18 66 419 – 1,928 2.32 – 9.09 (207)
Trachichthyidae Hoplostethus atlanticus ott816600 P 9 1.01 0.81 1.21 166 9,776 – 93,849 661.58 – 2509.01 (208)
Trichiuridae Aphanopus carbo ott113465 P 246 1.18 0.71 1.66 14 228,950 – 961,440 1950.46 – 4169.05 (209)
Zoarcidae Gymnelus viridis ott729337 D 248 1.26 0.8 1.74 1 11 13.80 (79)
Zoarcidae Lycodes esmarkii ott841648 D 249 0.93 0.38 1.48 1 1,200 4659.64 (79)
Zoarcidae Lycodes eudipleurostictus ott905435 D 250 1.23 0.65 1.8 1 186.5 183.60 (79)
Zoarcidae Lycodes frigidus ott539690 D 251 1.09 0.49 1.68 1 500 1207.55 (79)
Zoarcidae Lycodes pallidus ott178074 D 252 1.23 0.71 1.76 1 60 50.90 (79)
Zoarcidae Lycodes vahlii ott178063 D 255 1.04 0.59 1.49 1 37.5 7.04 (79)
Zoarcidae Melanostigma atlanticum ott3633033 D 256 0.94 0.52 1.35 3 36 – 56 3.94 – 7.54 (210)
* This is original data collected by the second author (D. Ross Robertson). Fish size were measured in standard length (mm), and total volume was measured as described in ref. (244). Then, for each female, clutch dry weight was obtained, and a
subsample of 50 eggs was obtained to estimate individual egg weight. Fecundity was estimated as the ratio between clutch dry weight and average individual egg weight.
16
Table S4. Summary of data used for Egg-volume–Mass model. This table shows summary stats of raw data used in Egg-volume–Mass model (Fig. 2B). OTL
= Open Tree of Life Identifier; SM = Spawning Mode (D = demersal, P = pelagic, S = scatterer, BM/BP/BI = mouth/pouch/internal brooder – all brooders have no
parental provisioning post fertilization); b1 / 2.5% / 97.5% = Estimates (mean and 95% Bayesian credible intervals) of species-specific egg-volume mass-scaling
exponents as obtained from Egg-volume–Mass model (see Materials and Methods); n = number of independent observations for each species.
Family Species OTL SM F3 b1 2.5% C.I. 97.5% C.I. n Egg-volume range (mm3) Mass range (g) Refs
Agonidae Leptagonus decagonus ott3632024 D 269 0.17 -0.23 0.55 1 4.189 32.32 (79)
Apogonidae Cheilodipterus macrodon ott965176 BM 59 0.21 -0.25 0.66 1 0.113 110.09 (80)
Apogonidae Holapogon maximus ott3635086 BM 61 0.15 -0.29 0.57 1 0.180 48.48 (80)
Apogonidae Jaydia hungi ott3634955 BM 62 0.08 -0.31 0.46 1 0.268 20.36 (80)
Apogonidae Ostorhinchus cookii ott102070 BM 63 0.07 -0.3 0.43 1 0.268 15.91 (80)
Apogonidae Ostorhinchus cyanosoma ott638185 BM 64 0.18 -0.14 0.49 1 0.113 6.02 (80)
Apogonidae Paroncheilus affinis ott974576 BM 66 0.33 -0.02 0.67 1 0.034 8.39 (211)
Apogonidae Siphamia tubifer ott5860863 BM 67 -0.09 -0.35 0.15 1 0.905 0.29 (80)
Apogonidae Taeniamia lineolata ott775407 BM 68 -0.04 -0.34 0.26 1 0.524 3.35 (80)
Atherinidae Atherina presbyter ott669918 D 2 0.12 -0.26 0.5 1 0.839 7.47 (81)
Atherinopsidae Menidia menidia ott189767 D 5 0.07 -0.16 0.29 20 0.624 – 0.860 3.48 – 14.03 (212)
Balistidae Balistes capriscus ott135140 S 335 0.18 -0.32 0.67 1 0.039 378.39 (85)
Bathydraconidae Akarotaxis nudiceps ott682053 D 69 -0.03 -0.4 0.33 1 9.203 16.00 (213)
Blenniidae Aidablennius sphynx ott898527 D 70 0.06 -0.28 0.39 1 0.195 0.63 (86)
Bothidae Arnoglossus laterna ott696263 P 258 0.27 -0.05 0.61 1 0.056 5.27 (87)
Callionymidae Synchiropus sechellensis ott3632718 P 73 0.23 -0.16 0.63 1 0.268 10.06 (214)
Carangidae Decapterus punctatus ott705034 P 74 0.41 0.01 0.82 1 0.024 31.62 (88)
Carangidae Trachurus mediterraneus ott174577 P 76 0.14 -0.33 0.6 1 0.271 364.39 (215)
Carangidae Trachurus picturatus ott174574 P 77 0.1 -0.25 0.46 8 0.194 – 0.194 38.10 – 73.11 (90)
Carangidae Trachurus trachurus ott472429 P 78 0.16 -0.21 0.54 3 0.342 – 0.435 140.72 – 317.47 (216); (215)
Channichthyidae Pagetopsis macropterus ott708993 D 84 0.01 -0.52 0.53 1 33.510 273.22 (217)
Clupeidae Alosa mediocris ott491656 D 10 0.14 -0.33 0.65 1 1.732 664.33 (218)
Clupeidae Amblygaster sirm ott328216 P 11 0.32 -0.11 0.77 1 0.048 42.93 (92)
Clupeidae Brevoortia patronus ott107260 P 12 0.08 -0.36 0.52 2 0.905 – 1.124 119.37 – 173.32 (219)
Clupeidae Ethmalosa fimbriata ott658559 P 16 0.25 -0.32 0.81 1 0.039 362.80 (94)
Clupeidae Nematalosa vlaminghi ott328199 P 17 0.07 -0.39 0.51 1 0.748 111.79 (95)
Clupeidae Spratelloides gracilis ott39732 D 21 0.12 -0.16 0.38 1 0.180 0.59 (99)
Congiopodidae Zanclorhynchus spinifer ott255677 P 271 0.25 -0.2 0.71 1 0.268 104.29 (213)
Cottidae Artediellus atlanticus ott654509 D 273 -0.12 -0.53 0.25 1 33.510 23.32 (79)
Cottidae Enophrys bison ott290389 D 274 0.25 -0.22 0.72 2 2.572 – 4.189 197.53 – 241.50 (101)
Cottidae Gymnocanthus tricuspis ott654474 D 275 0.19 -0.28 0.65 1 3.764 179.34 (79)
Cottidae Icelus bicornis ott3632116 D 276 0.12 -0.26 0.51 1 8.181 14.79 (79)
Cottidae Triglops pingelii ott568671 D 277 -0.01 -0.39 0.35 1 14.137 22.25 (79)
Cyclopteridae Cyclopteropsis mcalpini ott3631993 D 279 -0.07 -0.38 0.23 1 65.450 2.80 (79)
Cyclopteridae Eumicrotremus spinosus ott1097394 D 280 0.09 -0.34 0.52 1 29.764 60.65 (79)
Engraulidae Cetengraulis mysticetus ott1064953 P 24 0.2 0.04 0.36 86 0.048 – 0.113 25.10 – 71.50 (103)
Engraulidae Engraulis anchoita ott648864 P 25 0.12 -0.1 0.35 25 0.796 – 1.232 9.16 – 28.94 (220)
17
Table S4 Continued.
Family Species OTL SM F3 b1 2.5% C.I. 97.5% C.I. n Egg-volume range (mm3) Mass range (g) Refs
Engraulidae Engraulis mordax ott29754 P 26 -0.02 -0.37 0.32 1 1.023 14.74 (105)
Exocoetidae Hirundichthys affinis ott837204 D 7 0.04 -0.47 0.54 1 8.181 255.62 (108)
Fundulidae Fundulus heteroclitus ott4134765 D 29 0.15 -0.04 0.35 26 2.310 – 5.806 3.36 – 14.51 (221)
Fundulidae Fundulus xenicus ott839523 D 30 0.11 -0.19 0.4 1 3.131 1.75 (222)
Gadidae Boreogadus saida ott877055 P 33 -0.06 -0.43 0.31 1 2.550 18.48 (79)
Gadidae Eleginus nawaga ott450082 D 34 -0.02 -0.46 0.4 1 2.572 74.86 (79)
Gadidae Gadus morhua ott5860467 P 35 0.12 0.03 0.2 67 0.975 – 2.040 1382.51 – 24086.31 (110)
Gadidae Melanogrammus aeglefinus ott635904 P 36 0.08 0.02 0.14 142 1.018 – 1.736 111.75 – 2235.27 (223)
Gadidae Micromesistius australis ott1091056 P 38 0.1 -0.43 0.59 2 0.211 – 0.211 891.18 – 891.18 (114)
Gasterosteidae Gasterosteus aculeatus ott111122 D 45 0.24 -0.07 0.54 1 1.732 1.11 (116)
Gobiidae Elacatinus oceanops ott638369 D 89 -0.1 -0.36 0.15 1 0.776 0.25 (117)
Gobiidae Gobiosoma robustum ott1064480 D 90 0.18 -0.31 0.65 1 0.130 158.08 (118)
Gobiidae Oligolepis acutipennis ott767535 D 91 0.25 -0.21 0.7 1 0.022 25.48 (224)
Gobiidae Paragobiodon echinocephalus ott940827 D 92 0.1 -0.17 0.37 1 0.144 0.23 (119)
Gobiidae Paragobiodon lacunicolus ott758973 D 93 0.1 -0.18 0.37 1 0.144 0.23 (119)
Gobiidae Paragobiodon xanthosoma ott770378 D 94 0.1 -0.17 0.37 1 0.144 0.23 (119)
Haemulidae Haemulopsis corvinaeformis ott177515 P 97 0.21 -0.2 0.63 1 0.031 51.66 (108)
Hemiramphidae Hemiramphus brasiliensis ott953384 D 8 0.34 -0.13 0.83 1 0.268 111.47 (108)
Hemitripteridae Hemitripterus americanus ott160287 D 282 1.34 0.88 1.77 6 0.641 – 36.087 303.44 – 1004.16 (121)
Hexagrammidae Oxylebius pictus ott131554 D 284 0.3 -0.1 0.72 2 1.150 – 1.437 46.13 – 87.76 (122)
Labridae Anampses caeruleopunctatus ott372276 D 98 0.15 -0.32 0.63 1 0.134 331.12 (225)
Labridae Anampses twistii ott762167 P 99 0.12 -0.31 0.54 1 0.151 85.50 (225)
Labridae Cheilinus fasciatus ott155291 P 103 0.14 -0.33 0.63 1 0.137 637.55 (225)
Labridae Cheilinus oxycephalus ott195575 P 104 0.14 -0.3 0.59 1 0.125 114.33 (225)
Labridae Cheilinus trilobatus ott924675 P 105 0.12 -0.35 0.58 1 0.131 1556.96 (225)
Labridae Coris gaimard ott1092680 P 107 0.15 -0.33 0.62 1 0.134 294.30 (225)
Labridae Coris variegata ott3636534 P 108 0.15 -0.28 0.58 1 0.113 82.58 (225)
Labridae Epibulus insidiator ott120301 P 110 0.14 -0.35 0.61 1 0.154 1215.59 (225)
Labridae Gomphosus varius ott358606 P 111 0.18 -0.3 0.65 1 0.090 239.62 (225)
Labridae Halichoeres biocellatus ott1015700 P 112 0.12 -0.23 0.46 1 0.122 14.76 (225)
Labridae Halichoeres hortulanus ott694390 P 118 0.16 -0.31 0.63 1 0.113 228.47 (225)
Labridae Halichoeres melanurus ott627330 P 122 0.15 -0.29 0.59 1 0.131 53.47 (225)
Labridae Hemigymnus fasciatus ott924674 P 130 0.14 -0.3 0.58 1 0.195 2850.00 (225)
Labridae Labrichthys unilineatus ott302371 P 132 0.11 -0.29 0.51 1 0.168 50.73 (225)
Labridae Labroides dimidiatus ott302377 P 133 0.11 -0.23 0.46 1 0.137 17.95 (225)
Labridae Labropsis micronesica ott3636703 P 134 0.01 -0.32 0.34 1 0.258 10.96 (225)
Labridae Labropsis xanthonota ott3636702 P 135 0.08 -0.25 0.42 1 0.165 13.01 (225)
Labridae Macropharyngodon meleagris ott552117 P 136 0.1 -0.25 0.46 1 0.141 20.06 (225)
Labridae Novaculichthys taeniourus ott456983 P 137 0.15 -0.32 0.63 1 0.108 413.18 (225)
18
Table S4 Continued.
Family Species OTL SM F3 b1 2.5% C.I. 97.5% C.I. n Egg-volume range (mm3) Mass range (g) Refs
Labridae Oxycheilinus unifasciatus ott352271 P 138 0.14 -0.33 0.6 1 0.151 263.59 (225)
Labridae Pseudocheilinus hexataenia ott712600 P 139 0.14 -0.25 0.53 1 0.113 36.29 (225)
Labridae Symphodus roissali ott876619 D 142 0.06 -0.11 0.23 10 0.180 – 0.225 4.11 – 22.47 (226)
Liparidae Careproctus reinhardti ott1065176 D 285 0.12 -0.36 0.6 1 40.765 357.25 (79)
Liparidae Liparis fabricii ott496423 D 286 0.22 -0.22 0.66 1 7.148 60.36 (79)
Liparidae Paraliparis bathybius ott309222 BM 287 0.08 -0.37 0.53 1 36.087 122.99 (79)
Lutjanidae Lutjanus carponotatus ott333348 P 152 0.15 -0.15 0.45 4 0.032 – 0.042 90.62 – 431.79 (125)
Malacanthidae Caulolatilus microps ott39883 P 156 0.21 -0.25 0.67 1 0.258 2274.90 (129)
Merlucciidae Merluccius merluccius ott5860466 P 44 0.24 0.15 0.35 209 0.258 – 0.963 626.18 – 3465.51 (132)
Monacanthidae Brachaluteres jacksonianus ott333693 S 336 -0.02 -0.29 0.24 1 0.212 1.24 (133)
Monacanthidae Eubalichthys bucephalus ott3631310 S 337 0.13 -0.35 0.6 1 0.204 434.98 (134)
Monacanthidae Oxymonacanthus longirostris ott333706 S 338 0.01 -0.3 0.31 1 0.180 3.62 (135)
Monacanthidae Paramonacanthus japonicus ott3631319 S 339 0.13 -0.2 0.46 1 0.078 7.40 (136)
Monacanthidae Rudarius ercodes ott511593 S 340 0.11 -0.18 0.4 1 0.078 2.98 (136)
Mugilidae Chelon labrosus ott777079 P 46 -0.05 -0.46 0.33 13 0.090 – 0.410 1240.00 – 2300.00 (138)
Mugilidae Chelon ramada ott668241 P 47 0.08 -0.39 0.52 2 0.050 – 0.050 850.00 – 1450.00 (138)
Mugilidae Mugil cephalus ott418648 P 48 -0.11 -0.51 0.27 2 0.082 – 0.144 702.39 – 1833.16 (141); (139)
Mugilidae Planiliza subviridis ott823526 P 53 0.04 -0.32 0.41 1 0.157 17.13 (143)
Nototheniidae Lepidonotothen nudifrons ott260052 D 160 0.15 -0.31 0.6 1 2.806 117.20 (144)
Nototheniidae Patagonotothen longipes ott430554 D 161 0.33 -0.03 0.71 1 0.524 17.05 (227)
Nototheniidae Patagonotothen sima ott874773 D 162 0.46 0.17 0.76 1 0.128 2.43 (227)
Paralichthyidae Paralichthys dentatus ott1019735 P 259 0.16 -0.25 0.56 1 0.524 7935.57 (146)
Pleuronectidae Ammotretis rostratus ott600529 P 261 0.11 -0.33 0.56 4 0.508 – 0.572 356.19 – 563.87 (228)
Pleuronectidae Pseudopleuronectes americanus ott261419 D 265 0.07 -0.16 0.31 17 0.221 – 0.333 332.81 – 1322.71 (152)
Pleuronectidae Rhombosolea tapirina ott867269 P 267 0.11 -0.3 0.51 5 0.221 – 0.268 272.82 – 553.92 (228)
Pomacentridae Abudefduf saxatilis ott405751 D 167 0.02 -0.38 0.41 1 0.369 22.12 (155)
Pomacentridae Pomacentrus coelestis ott622060 D 180 0.08 -0.21 0.36 9 0.119 – 0.239 1.81 – 2.46 (157)
Pristigasteridae Ilisha africana ott741177 P 28 0.14 -0.26 0.53 1 0.322 37.69 (159)
Scaridae Cetoscarus bicolor ott1041720 P 190 0.14 -0.29 0.59 1 0.221 3384.72 (225)
Scaridae Chlorurus sordidus ott436737 P 191 0.14 -0.34 0.62 1 0.169 442.00 (225)
Scaridae Hipposcarus longiceps ott465039 P 193 0.16 -0.31 0.62 1 0.257 1527.80 (225)
Scaridae Scarus forsteni ott548869 P 195 0.14 -0.33 0.62 1 0.229 1077.09 (225)
Scaridae Scarus frenatus ott436725 P 196 0.13 -0.34 0.59 1 0.173 1851.12 (225)
Scaridae Scarus globiceps ott514869 P 197 0.11 -0.36 0.58 1 0.241 272.43 (225)
Scaridae Scarus schlegeli ott514881 P 201 0.12 -0.34 0.56 1 0.167 2441.14 (225)
Sciaenidae Bairdiella chrysoura ott761907 P 207 0.05 -0.38 0.47 1 0.172 54.91 (160)
Sciaenidae Cynoscion regalis ott5860595 P 208 0.13 -0.35 0.59 1 0.113 2018.26 (161)
Sciaenidae Larimus fasciatus ott897432 P 213 0.18 -0.21 0.57 1 0.058 25.23 (164)
Scombridae Auxis rochei ott713972 P 221 0.13 -0.35 0.62 1 0.362 453.17 (169)
19
Table S4 Continued.
Family Species OTL SM F3 b1 2.5% C.I. 97.5% C.I. n Egg-volume range (mm3) Mass range (g) Refs
Scombridae Euthynnus affinis ott308760 P 222 0.05 -0.35 0.45 3 0.123 – 0.191 5941.53 – 7549.52 (229)
Scombridae Katsuwonus pelamis ott308754 P 223 0.23 -0.11 0.58 10 0.144 – 0.248 2957.32 – 6503.90 (229)
Scombridae Sarda sarda ott833175 P 224 0.23 -0.24 0.7 1 1.124 1985.06 (230)
Scombridae Scomberomorus cavalla ott70546 P 226 0.14 -0.27 0.55 1 0.456 6702.24 (171)
Scombridae Thunnus albacares ott833188 P 229 0.09 -0.13 0.3 5 0.144 – 0.197 4620.11 – 43292.47 (229); (175)
Scombridae Thunnus obesus ott358613 P 230 0.17 -0.15 0.49 1 0.615 36847.75 (231)
Scombridae Thunnus thynnus ott358607 P 231 -0.17 -0.68 0.27 6 0.074 – 0.697 1690.13 – 2103.10 (232); (176)
Sebastidae Sebastes mentella ott1088432 BI 309 0.14 -0.36 0.62 1 0.685 690.02 (190)
Sebastidae Sebastes norvegicus ott1088441 BI 312 0.12 -0.37 0.61 1 0.685 1153.76 (190)
Sebastidae Sebastolobus altivelis ott811927 P 324 0.14 -0.35 0.61 1 1.023 288.33 (233)
Serranidae Cephalopholis cruentata ott12880 P 232 0.14 -0.33 0.63 1 0.078 220.12 (196)
Serranidae Epinephelus tauvina ott218649 P 236 0.05 -0.43 0.51 1 0.048 3094.02 (198)
Siganidae Siganus canaliculatus ott199059 S 241 0.19 -0.29 0.68 1 0.018 245.10 (200)
Soleidae Solea solea ott710233 P 268 0.13 -0.16 0.43 23 1.202 – 1.583 17000.00 – 51000.00 (234)
Sparidae Acanthopagrus latus ott760733 P 242 0.14 -0.35 0.63 1 0.100 570.93 (202)
Sparidae Archosargus rhomboidalis ott724892 P 243 0.3 -0.14 0.74 1 0.014 71.27 (203)
Sparidae Sparidentex hasta ott845300 P 244 0.16 -0.27 0.6 1 0.100 4986.75 (202)
Stromateidae Pampus chinensis ott916290 P 245 0.14 -0.34 0.63 1 0.606 741.00 (204)
Syngnathidae Entelurus aequoreus ott971459 BP 325 0.29 0.03 0.54 9 0.358 – 0.779 3.99 – 12.09 (235)
Syngnathidae Micrognathus crinitus ott3642321 BP 327 0.14 -0.09 0.35 1 0.796 0.17 (206)
Syngnathidae Nerophis ophidion ott806821 BP 328 0.17 -0.05 0.39 19 0.158 – 0.864 0.72 – 2.27 (235)
Syngnathidae Syngnathus acus ott774440 BP 329 0.21 0.02 0.4 19 6.118 – 11.706 12.97 – 62.60 (235)
Syngnathidae Syngnathus floridae ott774435 BP 330 0.37 0.07 0.66 1 0.322 3.03 (206)
Syngnathidae Syngnathus louisianae ott774414 BP 331 0.33 0 0.67 1 0.524 8.36 (206)
Syngnathidae Syngnathus rostellatus ott774411 BP 332 0.3 0.12 0.49 18 0.398 – 1.327 0.39 – 1.10 (235)
Syngnathidae Syngnathus scovelli ott251400 BP 333 0.22 -0.04 0.46 1 0.796 0.88 (206)
Syngnathidae Syngnathus typhle ott251396 BP 334 0.18 0.02 0.33 25 2.031 – 5.610 0.96 – 6.44 (235)
Tetraodontidae Canthigaster valentini ott701127 S 342 0.01 -0.33 0.34 1 0.180 4.18 (207)
Trichiuridae Aphanopus carbo ott113465 P 246 0.18 -0.27 0.64 1 0.806 2814.83 (209)
Zoarcidae Gymnelus viridis ott729337 D 248 0.11 -0.24 0.44 1 52.645 13.80 (79)
Zoarcidae Lycodes esmarkii ott841648 D 249 0.16 -0.25 0.57 1 113.097 4659.64 (79)
Zoarcidae Lycodes eudipleurostictus ott905435 D 250 0.13 -0.34 0.58 1 75.766 183.60 (79)
Zoarcidae Lycodes frigidus ott539690 D 251 0.18 -0.29 0.65 1 179.594 1207.55 (79)
Zoarcidae Lycodes pallidus ott178074 D 252 0.25 -0.16 0.65 1 22.449 50.90 (79)
Zoarcidae Lycodes polaris ott34255 D 253 0.16 -0.27 0.58 1 47.713 73.29 (79)
Zoarcidae Lycodes reticulatus ott752411 D 254 0.04 -0.46 0.54 1 344.791 271.32 (79)
Zoarcidae Lycodes vahlii ott178063 D 255 0.17 -0.15 0.47 1 33.510 7.04 (79)
Zoarcidae Melanostigma atlanticum ott3633033 D 256 0.22 -0.08 0.5 3 11.494 – 20.580 3.94 – 7.54 (210)
Zoarcidae Pachycara brachycephalum ott203179 D 257 0.12 -0.35 0.58 1 65.450 150.21 (213)
20
Table S5. Summary of data used for Egg-energy–Volume model. This table shows summary stats of raw data used in Egg-energy–Volume model (Fig. 2C).
OTL = Open Tree of Life Identifier; SM = Spawning Mode (D = demersal, P = pelagic, S = scatterer, BM/BP/BI = mouth/pouch/internal brooder – all brood-
ers have no parental provisioning post fertilization); b1 / 2.5% / 97.5% = Estimates (mean and 95% Bayesian credible intervals) of species-specific egg-energy
volume-scaling exponents as obtained from Egg-energy–Volume model (see Materials and Methods); n = number of independent measurements for each species.
Family Species OTL SM F3 b1 2.5% C.I. 97.5% C.I. n Egg-energy range (J) Egg-volume range (mm3) Refs
Acanthuridae Acanthurus coeruleus ott324283 P 55 0.76 0.34 1.19 6 0.24 – 0.29 0.136 – 0.163 *
Acanthuridae Acanthurus nigrofuscus ott605289 P 56 0.62 0.29 0.92 14 0.16 – 0.32 0.085 – 0.112 *
Acanthuridae Acanthurus triostegus ott467300 P 57 0.68 0.34 1.03 4 0.20 – 0.23 0.079 – 0.095 *
Acanthuridae Zebrasoma scopas ott199073 P 58 0.74 0.31 1.17 8 0.22 – 0.36 0.121 – 0.141 *
Agonidae Podothecus sachi ott590745 D 270 0.59 0.27 0.92 3 14.90 – 16.19 1.720 – 1.720 *
Apogonidae Cheilodipterus quinquelineatus ott364614 BM 60 0.61 0.2 1 2 0.96 – 1.06 0.298 – 0.377 *
Apogonidae Ostorhinchus cyanosoma ott638185 BM 64 0.71 0.31 1.1 5 0.49 – 0.67 0.105 – 0.140 *
Apogonidae Ostorhinchus doederleini ott687634 BM 65 0.78 0.31 1.26 1 1.16 0.233 *
Atherinopsidae Membras gilberti ott680137 D 4 0.72 0.29 1.15 5 3.70 – 4.20 0.852 – 1.066 *
Blenniidae Ophioblennius atlanticus ott282319 D 71 0.87 0.49 1.27 28 0.62 – 0.94 0.124 – 0.154 *
Blenniidae Ophioblennius steindachneri ott155044 D 72 0.84 0.51 1.17 24 0.38 – 0.80 0.102 – 0.140 *
Chaenopsidae Acanthemblemaria hancocki ott57340 D 80 0.63 0.27 0.97 42 1.34 – 2.44 0.303 – 0.403 *
Chaenopsidae Acanthemblemaria rivasi ott57343 D 81 0.74 0.48 0.99 36 2.28 – 3.48 0.477 – 0.696 *
Chaenopsidae Neoclinus uninotatus ott3639321 D 82 0.77 0.45 1.09 1 5.41 1.345 *
Chaetodontidae Chaetodon capistratus ott264060 P 83 0.71 0.25 1.15 8 0.29 – 0.43 0.154 – 0.184 *
Clupeidae Brevoortia tyrannus ott107265 P 13 0.39 0.1 0.67 1 0.98 2.456 *
Clupeidae Clupea pallasii ott9666 D 15 0.97 0.64 1.29 4 4.87 – 6.04 1.564 – 1.667 *
Coryphaenidae Coryphaena hippurus ott223663 P 85 0.71 0.34 1.06 1 4.17 2.600 *
Cottidae Alcichthys elongatus ott3632242 D 272 0.69 0.31 1.05 4 5.69 – 7.09 1.420 – 1.420 *
Cyclopteridae Aptocyclus ventricosus ott989887 D 278 1.03 0.68 1.37 1 46.35 2.430 *
Engraulidae Engraulis mordax ott29754 P 26 0.84 0.58 1.11 2 0.43 – 1.24 0.172 – 0.520 (236)
Engraulidae Engraulis ringens ott211981 P 27 0.91 0.49 1.34 2 0.86 – 0.98 0.284 – 0.322 (237)
Fundulidae Lucania goodei ott84261 D 31 0.78 0.46 1.1 1 7.50 1.193 (238)
Fundulidae Lucania parva ott839518 D 32 0.79 0.41 1.17 1 5.77 0.828 (238)
Gadidae Gadus morhua ott5860467 P 35 0.87 0.72 1.03 16 2.05 – 3.30 1.033 – 1.640 (239)
Gadidae Melanogrammus aeglefinus ott635904 P 36 0.87 0.74 1.01 60 2.17 – 3.72 1.166 – 1.957 (239)
Gadidae Merlangius merlangus ott635911 P 37 0.79 0.61 0.98 40 1.07 – 1.95 0.587 – 0.998 (239)
Gadidae Pollachius pollachius ott1014047 P 39 0.83 0.62 1.04 14 1.50 – 1.73 0.728 – 0.860 (239)
Gadidae Pollachius virens ott62649 P 40 0.87 0.65 1.1 8 1.43 – 1.85 0.656 – 0.790 (239)
Gadidae Trisopterus esmarkii ott926146 P 41 0.82 0.59 1.04 31 1.20 – 1.75 0.604 – 0.891 (239)
Gasterosteidae Gasterosteus aculeatus ott111122 D 45 0.4 0.05 0.74 3 2.57 – 2.91 2.915 – 3.246 (240)
Gobiidae Coryphopterus dicrus ott37549 D 86 0.62 0.29 0.94 11 0.27 – 0.33 0.082 – 0.108 *
Gobiidae Coryphopterus glaucofraenum ott494031 D 87 0.95 0.69 1.22 20 0.10 – 0.31 0.064 – 0.093 *
Gobiidae Coryphopterus personatus ott740192 D 88 0.77 0.51 1.02 28 0.13 – 0.31 0.059 – 0.082 *
Haemulidae Haemulon flavolineatum ott432196 P 96 0.73 0.28 1.17 5 0.56 – 0.63 0.318 – 0.350 *
Hemitripteridae Blepsias cirrhosus ott446859 D 281 1.1 0.83 1.38 1 114.86 3.170 *
Hemitripteridae Hemitripterus villosus ott438425 D 283 1.27 1.03 1.51 4 271.44 – 299.41 4.520 – 4.520 *
Hexagrammidae Oxylebius pictus ott131554 D 284 0.66 0.27 1.04 1 5.50 1.465 *
Labridae Bodianus axillaris ott548889 P 100 0.76 0.31 1.23 2 0.32 – 0.33 0.195 – 0.204 *
21
Table S5 Continued.
Family Species OTL SM F3 b1 2.5% C.I. 97.5% C.I. n Egg-energy range (J) Egg-volume range (mm3) Refs
Labridae Bodianus diplotaenia ott919235 P 101 0.67 0.19 1.11 10 0.16 – 0.32 0.172 – 0.211 *
Labridae Bodianus rufus ott372258 P 102 0.82 0.37 1.29 4 0.30 – 0.45 0.179 – 0.218 *
Labridae Clepticus parrae ott948303 P 106 0.76 0.33 1.19 6 0.24 – 0.35 0.151 – 0.166 *
Labridae Doratonotus megalepis ott763958 P 109 0.84 0.43 1.26 9 0.10 – 0.15 0.080 – 0.098 *
Labridae Halichoeres bivittatus ott694396 P 113 0.8 0.47 1.13 17 0.12 – 0.22 0.094 – 0.129 *
Labridae Halichoeres chierchiae ott615862 P 114 0.64 0.21 1.06 15 0.15 – 0.31 0.125 – 0.166 *
Labridae Halichoeres chloropterus ott1015715 P 115 0.79 0.36 1.23 3 0.15 – 0.18 0.086 – 0.103 *
Labridae Halichoeres dispilus ott762170 P 116 0.63 0.18 1.04 15 0.07 – 0.19 0.093 – 0.115 *
Labridae Halichoeres garnoti ott665840 P 117 0.69 0.27 1.11 15 0.17 – 0.29 0.117 – 0.162 *
Labridae Halichoeres hortulanus ott694390 P 118 0.94 0.57 1.31 2 0.13 – 0.14 0.095 – 0.101 *
Labridae Halichoeres maculipinna ott955368 P 119 0.78 0.34 1.22 6 0.13 – 0.18 0.090 – 0.107 *
Labridae Halichoeres margaritaceus ott694405 P 120 0.74 0.46 1.01 5 0.13 – 0.18 0.077 – 0.092 *
Labridae Halichoeres marginatus ott694408 P 121 0.55 0.19 0.9 3 0.20 – 0.24 0.096 – 0.105 *
Labridae Halichoeres melanurus ott627330 P 122 0.79 0.33 1.24 6 0.19 – 0.22 0.111 – 0.114 *
Labridae Halichoeres miniatus ott694399 P 123 0.81 0.45 1.16 3 0.15 – 0.19 0.097 – 0.114 *
Labridae Halichoeres nebulosus ott1015709 P 124 0.78 0.34 1.21 1 0.18 0.093 *
Labridae Halichoeres nicholsi ott694402 P 125 0.71 0.28 1.13 2 0.20 – 0.25 0.137 – 0.138 *
Labridae Halichoeres notospilus ott694414 P 126 0.96 0.64 1.28 3 0.13 – 0.14 0.094 – 0.099 *
Labridae Halichoeres pictus ott173446 P 127 1 0.57 1.46 4 0.16 – 0.43 0.115 – 0.154 *
Labridae Halichoeres poeyi ott38828 P 128 0.68 0.24 1.1 15 0.19 – 0.28 0.117 – 0.143 *
Labridae Halichoeres trimaculatus ott942026 P 129 0.8 0.35 1.25 6 0.14 – 0.18 0.088 – 0.099 *
Labridae Hemigymnus fasciatus ott924674 P 130 0.75 0.29 1.22 1 0.37 0.181 *
Labridae Hemigymnus melapterus ott302368 P 131 0.67 0.25 1.1 6 0.30 – 0.43 0.143 – 0.186 *
Labridae Novaculichthys taeniourus ott456983 P 137 0.72 0.3 1.13 6 0.12 – 0.21 0.085 – 0.101 *
Labridae Stethojulis bandanensis ott201174 P 140 0.79 0.49 1.08 2 0.12 – 0.12 0.069 – 0.071 *
Labridae Symphodus mediterraneus ott730049 D 141 0.78 0.33 1.22 1 1.05 0.236 *
Labridae Symphodus roissali ott876619 D 142 0.49 0.1 0.86 10 0.89 – 1.01 0.200 – 0.280 *
Labridae Symphodus rostratus ott553190 D 143 0.75 0.37 1.12 6 1.09 – 1.23 0.266 – 0.305 *
Labridae Symphodus tinca ott787184 D 144 0.87 0.45 1.28 3 1.17 – 1.35 0.251 – 0.271 *
Labridae Thalassoma bifasciatum ott907850 P 145 0.94 0.64 1.23 15 0.08 – 0.15 0.072 – 0.092 *
Labridae Thalassoma hardwicke ott102998 P 146 0.69 0.43 0.95 7 0.11 – 0.14 0.058 – 0.068 *
Labridae Thalassoma jansenii ott1094748 P 147 0.77 0.51 1.02 6 0.11 – 0.13 0.060 – 0.075 *
Labridae Thalassoma lucasanum ott1094750 P 148 0.78 0.5 1.05 15 0.09 – 0.16 0.065 – 0.083 *
Labridae Thalassoma lunare ott948286 P 149 0.71 0.44 0.97 5 0.12 – 0.13 0.065 – 0.074 *
Moronidae Dicentrarchus labrax ott3549 P 157 0.81 0.5 1.13 17 2.35 – 3.67 0.776 – 1.232 (241)
Mullidae Parupeneus multifasciatus ott295485 P 158 0.76 0.28 1.23 4 0.26 – 0.31 0.126 – 0.137 *
Muraenesocidae Muraenesox cinereus ott479856 P 1 0.59 0.35 0.82 7 3.20 – 5.79 2.070 – 6.290 *
Nemipteridae Scolopsis bilineata ott463069 P 159 0.76 0.29 1.22 1 0.45 0.227 *
Pinguipedidae Parapercis snyderi ott3635246 P 163 0.67 0.25 1.08 6 0.25 – 0.35 0.131 – 0.203 *
Pleuronectidae Hippoglossus hippoglossus ott408328 P 263 0.84 0.74 0.95 6 37.25 – 40.75 15.150 – 16.056 (242)
Pleuronectidae Pleuronectes platessa ott408337 P 264 0.68 0.56 0.81 41 5.22 – 7.96 2.802 – 4.489 (151)
Pleuronectidae Pseudopleuronectes americanus ott261419 D 265 0.5 0.14 0.85 2 1.25 – 1.28 0.230 – 0.463 *
Pomacentridae Abudefduf bengalensis ott318931 D 166 0.92 0.51 1.32 6 1.23 – 1.59 0.227 – 0.262 *
22
Table S5 Continued.
Family Species OTL SM F3 b1 2.5% C.I. 97.5% C.I. n Egg-energy range (J) Egg-volume range (mm3) Refs
Pomacentridae Abudefduf saxatilis ott405751 D 167 0.67 0.27 1.05 48 0.87 – 1.17 0.171 – 0.214 *
Pomacentridae Abudefduf septemfasciatus ott129790 D 168 0.77 0.36 1.18 10 1.06 – 1.29 0.221 – 0.237 *
Pomacentridae Abudefduf sordidus ott1053071 D 169 0.79 0.42 1.17 8 1.07 – 1.47 0.233 – 0.269 *
Pomacentridae Abudefduf troschelii ott961357 D 170 0.87 0.54 1.21 44 0.57 – 1.13 0.133 – 0.172 *
Pomacentridae Abudefduf vaigiensis ott1053067 D 171 0.76 0.37 1.16 13 0.76 – 1.01 0.137 – 0.178 *
Pomacentridae Chromis atrilobata ott436999 D 173 1.1 0.84 1.35 6 0.31 – 0.40 0.075 – 0.088 *
Pomacentridae Chromis atripectoralis ott741423 D 174 0.67 0.41 0.9 5 0.48 – 0.59 0.066 – 0.091 *
Pomacentridae Chromis multilineata ott437016 D 175 0.83 0.6 1.04 14 0.35 – 0.50 0.070 – 0.081 *
Pomacentridae Hypsypops rubicundus ott847666 D 176 0.84 0.59 1.07 7 1.54 – 1.93 0.401 – 0.453 *
Pomacentridae Microspathodon bairdii ott237630 D 177 0.73 0.33 1.09 10 0.57 – 0.78 0.133 – 0.175 *
Pomacentridae Microspathodon chrysurus ott847660 D 178 0.72 0.42 1.02 38 0.42 – 0.79 0.109 – 0.140 *
Pomacentridae Microspathodon dorsalis ott205758 D 179 0.77 0.5 1.04 24 0.51 – 0.87 0.115 – 0.196 *
Pomacentridae Stegastes acapulcoensis ott3635554 D 181 0.96 0.69 1.21 22 0.33 – 0.56 0.092 – 0.103 *
Pomacentridae Stegastes adustus ott323181 D 182 0.97 0.76 1.18 40 0.26 – 0.62 0.077 – 0.115 *
Pomacentridae Stegastes diencaeus ott729161 D 183 0.72 0.33 1.08 18 0.62 – 0.82 0.133 – 0.156 *
Pomacentridae Stegastes flavilatus ott3635541 D 184 0.9 0.69 1.09 27 0.21 – 0.58 0.069 – 0.118 *
Pomacentridae Stegastes leucostictus ott100830 D 186 0.89 0.48 1.27 41 1.03 – 1.45 0.213 – 0.261 *
Pomacentridae Stegastes partitus ott345269 D 187 0.94 0.77 1.13 40 0.19 – 0.54 0.061 – 0.088 *
Pomacentridae Stegastes planifrons ott665837 D 188 0.84 0.48 1.2 19 0.45 – 0.91 0.127 – 0.151 *
Pomacentridae Stegastes variabilis ott323173 D 189 0.96 0.59 1.33 24 0.45 – 0.84 0.133 – 0.172 *
Scaridae Chlorurus sordidus ott436737 P 191 0.67 0.26 1.05 6 0.18 – 0.29 0.099 – 0.121 *
Scaridae Cryptotomus roseus ott321483 P 192 0.97 0.61 1.33 10 0.10 – 0.16 0.092 – 0.110 *
Scaridae Scarus flavipectoralis ott1041716 P 194 0.71 0.31 1.11 2 0.22 – 0.23 0.122 – 0.124 *
Scaridae Scarus frenatus ott436725 P 196 0.68 0.27 1.07 2 0.25 – 0.27 0.121 – 0.123 *
Scaridae Scarus globiceps ott514869 P 197 0.79 0.35 1.23 1 0.24 0.137 *
Scaridae Scarus iseri ott686634 P 198 0.87 0.43 1.3 5 0.20 – 0.24 0.132 – 0.144 *
Scaridae Scarus niger ott14311 P 199 0.56 0.16 0.95 3 0.24 – 0.32 0.118 – 0.123 *
Scaridae Scarus psittacus ott514872 P 200 1.06 0.71 1.41 5 0.11 – 0.16 0.097 – 0.110 *
Scaridae Scarus schlegeli ott514881 P 201 0.64 0.22 1.05 4 0.23 – 0.27 0.125 – 0.137 *
Scaridae Scarus spinus ott419732 P 202 0.77 0.4 1.15 2 0.19 – 0.22 0.107 – 0.111 *
Scaridae Sparisoma atomarium ott686627 P 203 1.04 0.65 1.43 2 0.10 – 0.13 0.103 – 0.109 *
Scaridae Sparisoma aurofrenatum ott686624 P 204 0.57 0.15 0.96 5 0.19 – 0.29 0.117 – 0.130 *
Scaridae Sparisoma radians ott214109 P 205 0.78 0.47 1.08 6 0.10 – 0.18 0.090 – 0.109 *
Scaridae Sparisoma rubripinne ott686642 P 206 0.58 0.17 0.99 5 0.22 – 0.28 0.128 – 0.135 *
Sciaenidae Cynoscion regalis ott5860595 P 208 0.18 -0.22 0.54 10 0.62 – 2.38 0.349 – 0.534 *
Sciaenidae Leiostomus xanthurus ott1052160 P 214 0.62 0.12 1.07 1 0.61 0.276 *
Sciaenidae Micropogonias undulatus ott987393 P 217 0.69 0.18 1.16 1 0.68 0.220 *
Serranidae Epinephelus fasciatus ott534146 P 234 0.85 0.44 1.28 14 0.14 – 0.31 0.171 – 0.215 *
Serranidae Serranus tabacarius ott284266 P 238 0.79 0.39 1.18 4 0.39 – 0.47 0.331 – 0.343 *
Serranidae Serranus tigrinus ott807897 P 239 0.54 0.07 0.96 15 0.19 – 0.31 0.148 – 0.209 *
Serranidae Serranus tortugarum ott511782 P 240 0.75 0.32 1.19 15 0.22 – 0.44 0.200 – 0.248 *
Soleidae Solea solea ott710233 P 268 0.77 0.42 1.12 13 2.02 – 3.41 1.202 – 1.583 (234)
Tetraodontidae Canthigaster rostrata ott484093 D 341 0.73 0.26 1.19 1 0.69 0.122 *
Trichodontidae Arctoscopus japonicus ott883978 D 247 0.79 0.58 1 5 92.66 – 110.46 18.077 – 22.358 *
* This is original data collected by the second author (D. Ross Robertson). All remaining species data come from studies that measured egg dry weight rather than energy. Based on (27) and our own data (see Fig. S1), we assumed that egg energy
scaled isometrically with egg dry weight, with an average of ~25 J / mg of egg.
23
Table S6. Summary of data used for Total-volume–Mass model. OTL = Open Tree of Life Identifier; SM = Spawning Mode (D = demersal, P = pelagic, S =
scatterer, BM/BP/BI = mouth/pouch/internal brooder – all brooders have no parental provisioning post fertilization); F3 = Number ID in Fig. 3; β1 / 2.5% / 97.5%
= Estimates (mean and 95% Bayesian credible intervals) of species-specific total egg-volume mass-scaling exponents (see Materials and Methods); n = number of
independent observations for each species.
Family Species OTL SM F3 β1 2.5% C.I. 97.5% C.I. n Total egg-volume range (mm3) Mass range (g) Refs
Atherinidae Atherina presbyter ott669918 D 2 1.08 0.87 1.28 33 784.1 – 9,566 1.47 – 13.70 (81)
Balistidae Balistes capriscus ott135140 S 335 1.13 0.75 1.51 64 13,123 – 77,449 272.64 – 696.02 (85)
Bothidae Arnoglossus laterna ott696263 P 258 1.13 0.71 1.59 2 398 – 3,343 2.68 – 9.09 (87)
Carangidae Decapterus punctatus ott705034 P 74 1.48 0.99 2.01 2 151.5 – 1,246 16.71 – 53.59 (88)
Carangidae Trachurus picturatus ott174574 P 77 1.12 0.55 1.69 8 4,595 – 9,493 38.10 – 73.11 (90)
Clupeidae Amblygaster sirm ott328216 P 11 1.74 1.22 2.34 33 967.7 – 6,699 34.39 – 57.50 (92)
Clupeidae Ethmalosa fimbriata ott658559 P 16 0.91 0.51 1.3 32 613.5 – 2,013 182.55 – 582.47 (94)
Clupeidae Spratelloides gracilis ott39732 D 21 0.98 0.69 1.28 18 106.7 – 1,062 0.20 – 0.86 (99)
Cottidae Enophrys bison ott290389 D 274 1.14 0.4 1.91 2 48,362 – 133,622 197.53 – 241.50 (101)
Engraulidae Cetengraulis mysticetus ott1064953 P 24 1.38 1.11 1.66 86 642.1 – 6,957 25.10 – 71.50 (103)
Engraulidae Engraulis mordax ott29754 P 26 0.98 0.52 1.46 21 8,869 – 36,367 10.47 – 23.98 (105)
Gadidae Gadus morhua ott5860467 P 35 1.51 1.36 1.66 67 923,939 – 36,102,635 1382.51 – 24086.31 (110)
Gasterosteidae Gasterosteus aculeatus ott111122 D 45 0.91 0.51 1.31 2 330.8 – 905.9 0.80 – 1.50 (116)
Hemitripteridae Hemitripterus americanus ott160287 D 282 2.43 1.83 3.08 6 2,313 – 800,914 303.44 – 1004.16 (121)
Hexagrammidae Oxylebius pictus ott131554 D 284 0.73 0.46 1.01 43 14,712 – 42,473 32.22 – 148.53 (122)
Lutjanidae Lutjanus carponotatus ott333348 P 152 1.81 1.59 2.03 55 154 – 31,682 77.08 – 485.66 (125)
Malacanthidae Caulolatilus microps ott39883 P 156 1.5 1.22 1.79 43 21,548 – 1,063,744 762.87 – 4527.12 (129)
Merlucciidae Merluccius merluccius ott5860466 P 44 1.17 0.99 1.34 209 10,066 – 557,218 626.18 – 3465.51 (132)
Monacanthidae Eubalichthys bucephalus ott3631310 S 337 1.47 0.71 2.28 2 1,833 – 8,046 355.86 – 524.85 (134)
Monacanthidae Oxymonacanthus longirostris ott333706 S 338 1.35 0.89 1.81 3 17.96 – 35.92 3.30 – 3.79 (135)
Monacanthidae Rudarius ercodes ott511593 S 340 0.91 0.52 1.29 3 199.6 – 637.6 1.25 – 5.78 (136)
Mugilidae Chelon labrosus ott777079 P 46 1.01 0.33 1.66 13 31,770 – 180,570 1240.00 – 2300.00 (138)
Mugilidae Chelon ramada ott668241 P 47 1.23 0.49 1.98 2 29,050 – 62,150 850.00 – 1450.00 (138)
Mugilidae Mugil cephalus ott418648 P 48 0.95 0.74 1.16 73 37,102 – 395,750 274.75 – 3760.17 (139); (141)
Mugilidae Planiliza subviridis ott823526 P 53 1.02 0.56 1.5 12 6,011 – 22,974 11.69 – 28.41 (143)
Paralichthyidae Paralichthys dentatus ott1019735 P 259 1.06 0.9 1.22 134 240,886 – 2,200,444 2627.81 – 20166.83 (146)
Pleuronectidae Pseudopleuronectes americanus ott261419 D 265 0.99 0.58 1.4 17 64,023 – 416,296 332.81 – 1322.71 (152)
Pomacentridae Pomacentrus coelestis ott622060 D 180 0.97 0.68 1.26 81 268.2 – 934.8 1.27 – 3.26 (157)
Sciaenidae Cynoscion regalis ott5860595 P 208 1.51 1.24 1.79 29 10,181 – 244,728 554.68 – 3969.46 (161)
Sciaenidae Larimus fasciatus ott897432 P 213 1.04 0.86 1.22 81 1,021 – 18,297 8.75 – 59.86 (164)
Scombridae Scomberomorus cavalla ott70546 P 226 1.37 1.23 1.5 64 31,588 – 5,566,904 886.37 – 28991.83 (171)
Scombridae Thunnus thynnus ott358607 P 231 1.18 0.58 1.78 27 1,001,262 – 4,240,637 1336.98 – 2861.01 (176)
Sebastidae Sebastes mentella ott1088432 BI 309 1.44 1.14 1.75 54 7,152 – 79,378 381.88 – 1284.55 (190)
Sebastidae Sebastes norvegicus ott1088441 BI 312 1.02 0.63 1.41 26 24,234 – 146,044 477.49 – 2037.78 (190)
Serranidae Cephalopholis cruentata ott12880 P 232 1.22 0.65 1.79 12 6,720 – 47,971 142.95 – 414.81 (196)
Serranidae Epinephelus tauvina ott218649 P 236 0.81 0.33 1.28 4 40,565 – 138,602 1062.45 – 6012.89 (198)
Siganidae Siganus canaliculatus ott199059 S 241 1.27 1.05 1.48 27 649.8 – 28,482 59.67 – 870.20 (200)
Soleidae Solea solea ott710233 P 268 1.81 1.28 2.41 23 25,862 – 5,699,096 17000.00 – 51000.00 (234)
24
Table S6 Continued.
Family Species OTL SM F3 β1 2.5% C.I. 97.5% C.I. n Total egg-volume range (mm3) Mass range (g) Refs
Sparidae Acanthopagrus latus ott760733 P 242 1.11 0.35 1.87 5 135,589 – 214,311 426.21 – 660.47 (202)
Sparidae Archosargus rhomboidalis ott724892 P 243 0.95 0.71 1.19 23 250.7 – 2,210 15.05 – 156.17 (203)
Sparidae Sparidentex hasta ott845300 P 244 1.28 0.68 1.92 6 30,686 – 168,559 2604.06 – 6288.57 (202)
Stromateidae Pampus chinensis ott916290 P 245 1.17 0.41 1.93 8 48,733 – 56,431 647.18 – 876.77 (204)
Tetraodontidae Canthigaster valentini ott701127 S 342 0.98 0.76 1.2 66 75.25 – 346.3 2.32 – 9.09 (207)
Trichiuridae Aphanopus carbo ott113465 P 246 1.27 0.68 1.87 14 184,493 – 774,750 1950.46 – 4169.05 (209)
Zoarcidae Melanostigma atlanticum ott3633033 D 256 0.96 0.48 1.42 3 436.8 – 1,152 3.94 – 7.54 (210)
The above results were obtained with the subset of studies that contained paired measurements of fecundity (Table S3) and egg-volume (Table S4). This analysis was conducted by using the subset of studies that contained more than 1 observa-
tion.
25
Table S7. Species-specific reproductive-energy output mass-scaling exponents. This table contains all species present across the three main analyses in this
study (Fecundity–Mass, Egg-volume–Mass, and Egg-energy–Volume models). OTL = Open Tree of Life Identifier; SM = Spawning Mode (D = demersal, P =
pelagic, S = scatterer, BM/BP/BI = mouth/pouch/internal brooder – all brooders have no parental provisioning post fertilization); F3 = Number ID in Fig. 3; b1 /
2.5% / 97.5% = Estimates (mean and 95% Bayesian credible intervals) of reproductive-energy output mass-scaling exponents as obtained by combining posterior
distributions of Fecundity–Mass, Egg-volume–Mass, and Egg-energy–Volume models (see Materials and Methods); D1–3 = Is this species respectively present in
Datasets 1 (Fecundity), 2 (Egg-volume), and/or 3 (Egg-energy)?
Family Species OTL SM F3 b1 2.5% C.I. 97.5% C.I. D1 D2 D3
Acanthuridae Acanthurus coeruleus ott324283 P 55 1.29 1.19 1.41 No No Yes
Acanthuridae Acanthurus nigrofuscus ott605289 P 56 1.27 1.18 1.37 No No Yes
Acanthuridae Acanthurus triostegus ott467300 P 57 1.28 1.19 1.38 No No Yes
Acanthuridae Zebrasoma scopas ott199073 P 58 1.29 1.19 1.4 No No Yes
Agonidae Leptagonus decagonus ott3632024 D 269 1.3 0.71 1.87 Yes Yes No
Agonidae Podothecus sachi ott590745 D 270 1.27 1.18 1.36 No No Yes
Apogonidae Cheilodipterus macrodon ott965176 BM 59 1.44 0.78 2.11 Yes Yes No
Apogonidae Cheilodipterus quinquelineatus ott364614 BM 60 1.27 1.18 1.37 No No Yes
Apogonidae Holapogon maximus ott3635086 BM 61 1.36 0.72 1.95 Yes Yes No
Apogonidae Jaydia hungi ott3634955 BM 62 1.29 0.71 1.88 Yes Yes No
Apogonidae Ostorhinchus cookii ott102070 BM 63 1.32 0.75 1.87 Yes Yes No
Apogonidae Ostorhinchus cyanosoma* ott638185 BM 64 1.22 0.74 1.72 Yes Yes Yes
Apogonidae Ostorhinchus doederleini ott687634 BM 65 1.29 1.19 1.41 No No Yes
Apogonidae Paroncheilus affinis ott974576 BM 66 1.43 1.16 1.72 No Yes No
Apogonidae Siphamia tubifer ott5860863 BM 67 0.79 0.41 1.18 Yes Yes No
Apogonidae Taeniamia lineolata ott775407 BM 68 1.04 0.57 1.53 Yes Yes No
Atherinidae Atherina presbyter ott669918 D 2 1.19 0.84 1.53 Yes Yes No
Atherinopsidae Leuresthes tenuis ott274466 D 3 1.5 1.03 1.97 Yes No No
Atherinopsidae Membras gilberti ott680137 D 4 1.28 1.19 1.4 No No Yes
Atherinopsidae Menidia menidia ott189767 D 5 1.24 1.05 1.42 No Yes No
Atherinopsidae Odontesthes argentinensis ott93287 D 6 1.28 0.96 1.6 Yes No No
Balistidae Balistes capriscus ott135140 S 335 1.75 1.32 2.17 Yes Yes No
Bathydraconidae Akarotaxis nudiceps ott682053 D 69 1.16 0.88 1.44 No Yes No
Blenniidae Aidablennius sphynx ott898527 D 70 1.11 0.65 1.57 Yes Yes No
Blenniidae Ophioblennius atlanticus ott282319 D 71 1.31 1.2 1.42 No No Yes
Blenniidae Ophioblennius steindachneri ott155044 D 72 1.3 1.2 1.41 No No Yes
Bothidae Arnoglossus laterna ott696263 P 258 1.18 0.73 1.64 Yes Yes No
Callionymidae Synchiropus sechellensis ott3632718 P 73 1.36 1.05 1.67 No Yes No
Carangidae Decapterus punctatus ott705034 P 74 1.59 1.06 2.13 Yes Yes No
Carangidae Elagatis bipinnulata ott705026 P 75 1.1 0.83 1.37 Yes No No
Carangidae Trachurus mediterraneus ott174577 P 76 1.29 0.93 1.66 No Yes No
Carangidae Trachurus picturatus ott174574 P 77 1.21 0.65 1.77 Yes Yes No
Carangidae Trachurus trachurus ott472429 P 78 1.3 1.01 1.61 No Yes No
Centropomidae Centropomus undecimalis ott317368 P 79 1.6 1.19 2.01 Yes No No
Chaenopsidae Acanthemblemaria hancocki ott57340 D 80 1.27 1.18 1.37 No No Yes
Chaenopsidae Acanthemblemaria rivasi ott57343 D 81 1.29 1.2 1.38 No No Yes
26
Table S7 Continued.
Family Species OTL SM F3 b1 2.5% C.I. 97.5% C.I. D1 D2 D3
Clupeidae Amblygaster sirm ott328216 P 11 1.82 1.25 2.39 Yes Yes No
Clupeidae Brevoortia patronus ott107260 P 12 1.24 0.9 1.59 No Yes No
Clupeidae Brevoortia tyrannus ott107265 P 13 1.24 1.16 1.32 No No Yes
Clupeidae Clupea harengus ott1005932 D 14 1.65 1.52 1.78 Yes No No
Clupeidae Clupea pallasii ott9666 D 15 1.32 1.21 1.44 No No Yes
Clupeidae Ethmalosa fimbriata ott658559 P 16 1.09 0.55 1.64 Yes Yes No
Clupeidae Nematalosa vlaminghi ott328199 P 17 1.2 0.54 1.84 Yes Yes No
Clupeidae Opisthonema libertate ott437031 P 18 1.21 0.81 1.6 Yes No No
Clupeidae Opisthonema medirastre ott1195 P 19 1.35 0.92 1.78 Yes No No
Clupeidae Sardinops sagax ott121238 P 20 1.08 0.93 1.23 Yes No No
Clupeidae Spratelloides gracilis ott39732 D 21 1.08 0.76 1.42 Yes Yes No
Clupeidae Strangomera bentincki ott648871 P 22 1.19 0.75 1.6 Yes No No
Congiopodidae Zanclorhynchus spinifer ott255677 P 271 1.37 1.03 1.74 No Yes No
Coryphaenidae Coryphaena hippurus ott223663 P 85 1.28 1.19 1.39 No No Yes
Cottidae Alcichthys elongatus ott3632242 D 272 1.28 1.19 1.39 No No Yes
Cottidae Artediellus atlanticus ott654509 D 273 1.22 0.64 1.79 Yes Yes No
Cottidae Enophrys bison ott290389 D 274 1.32 0.64 2 Yes Yes No
Cottidae Gymnocanthus tricuspis ott654474 D 275 1.33 0.97 1.68 No Yes No
Cottidae Icelus bicornis ott3632116 D 276 1.19 0.63 1.75 Yes Yes No
Cottidae Triglops pingelii ott568671 D 277 1.11 0.54 1.67 Yes Yes No
Cyclopteridae Aptocyclus ventricosus ott989887 D 278 1.33 1.22 1.45 No No Yes
Cyclopteridae Cyclopteropsis mcalpini ott3631993 D 279 0.99 0.51 1.47 Yes Yes No
Cyclopteridae Eumicrotremus spinosus ott1097394 D 280 1.25 0.92 1.58 No Yes No
Engraulidae Anchoviella lepidentostole ott610213 P 23 1.08 0.7 1.46 Yes No No
Engraulidae Cetengraulis mysticetus ott1064953 P 24 1.39 1.11 1.66 Yes Yes No
Engraulidae Engraulis anchoita ott648864 P 25 1.36 1.13 1.58 Yes Yes No
Engraulidae Engraulis mordax* ott29754 P 26 1.15 0.64 1.64 Yes Yes Yes
Engraulidae Engraulis ringens ott211981 P 27 1.48 1.25 1.71 Yes No Yes
Exocoetidae Hirundichthys affinis ott837204 D 7 1.24 0.56 1.92 Yes Yes No
Fundulidae Fundulus heteroclitus ott4134765 D 29 1.3 1.15 1.46 No Yes No
Fundulidae Fundulus xenicus ott839523 D 30 1.26 1.02 1.5 No Yes No
Fundulidae Lucania goodei ott84261 D 31 1.29 1.2 1.4 No No Yes
Fundulidae Lucania parva ott839518 D 32 1.3 1.2 1.41 No No Yes
Gadidae Boreogadus saida ott877055 P 33 0.85 0.28 1.42 Yes Yes No
Gadidae Eleginus nawaga ott450082 D 34 1.1 0.47 1.73 Yes Yes No
Gadidae Gadus morhua* ott5860467 P 35 1.33 1.25 1.43 Yes Yes Yes
Gadidae Melanogrammus aeglefinus ott635904 P 36 1.25 1.17 1.34 No Yes Yes
Gadidae Merlangius merlangus ott635911 P 37 1.14 1.01 1.28 Yes No Yes
Gadidae Micromesistius australis ott1091056 P 38 1.64 1.19 2.07 Yes Yes No
Gadidae Pollachius pollachius ott1014047 P 39 1.3 1.21 1.4 No No Yes
Gadidae Pollachius virens ott62649 P 40 1.31 1.21 1.41 No No Yes
Gadidae Trisopterus esmarkii ott926146 P 41 1.3 1.2 1.4 No No Yes
Gasterosteidae Gasterosteus aculeatus* ott111122 D 45 1.01 0.82 1.23 Yes Yes Yes
Gobiidae Coryphopterus dicrus ott37549 D 86 1.27 1.18 1.37 No No Yes
27
Table S7 Continued.
Family Species OTL SM F3 b1 2.5% C.I. 97.5% C.I. D1 D2 D3
Gobiidae Coryphopterus glaucofraenum ott494031 D 87 1.32 1.21 1.42 No No Yes
Gobiidae Coryphopterus personatus ott740192 D 88 1.29 1.2 1.39 No No Yes
Gobiidae Elacatinus oceanops ott638369 D 89 0.72 0.3 1.12 Yes Yes No
Gobiidae Gobiosoma robustum ott1064480 D 90 1.49 0.78 2.17 Yes Yes No
Gobiidae Oligolepis acutipennis ott767535 D 91 1.37 1.02 1.74 No Yes No
Gobiidae Paragobiodon echinocephalus ott940827 D 92 0.95 0.53 1.37 Yes Yes No
Gobiidae Paragobiodon lacunicolus ott758973 D 93 0.96 0.54 1.37 Yes Yes No
Gobiidae Paragobiodon xanthosoma ott770378 D 94 1.04 0.62 1.45 Yes Yes No
Gobiidae Pomatoschistus minutus ott1048873 D 95 0.92 0.57 1.28 Yes No No
Haemulidae Haemulon flavolineatum ott432196 P 96 1.29 1.19 1.4 No No Yes
Haemulidae Haemulopsis corvinaeformis ott177515 P 97 0.69 0.3 1.08 Yes Yes No
Hemiramphidae Hemiramphus brasiliensis ott953384 D 8 1.1 0.55 1.67 Yes Yes No
Hemitripteridae Blepsias cirrhosus ott446859 D 281 1.34 1.23 1.45 No No Yes
Hemitripteridae Hemitripterus americanus ott160287 D 282 2.25 1.69 2.82 Yes Yes No
Hemitripteridae Hemitripterus villosus ott438425 D 283 1.36 1.24 1.49 No No Yes
Hexagrammidae Oxylebius pictus* ott131554 D 284 1.12 0.57 1.7 Yes Yes Yes
Labridae Anampses caeruleopunctatus ott372276 D 98 1.3 0.93 1.66 No Yes No
Labridae Anampses twistii ott762167 P 99 1.28 0.94 1.62 No Yes No
Labridae Bodianus axillaris ott548889 P 100 1.29 1.19 1.41 No No Yes
Labridae Bodianus diplotaenia ott919235 P 101 1.28 1.18 1.39 No No Yes
Labridae Bodianus rufus ott372258 P 102 1.3 1.19 1.42 No No Yes
Labridae Cheilinus fasciatus ott155291 P 103 1.29 0.92 1.67 No Yes No
Labridae Cheilinus oxycephalus ott195575 P 104 1.29 0.95 1.64 No Yes No
Labridae Cheilinus trilobatus ott924675 P 105 1.28 0.92 1.63 No Yes No
Labridae Clepticus parrae ott948303 P 106 1.29 1.19 1.4 No No Yes
Labridae Coris gaimard ott1092680 P 107 1.29 0.92 1.66 No Yes No
Labridae Coris variegata ott3636534 P 108 1.3 0.96 1.63 No Yes No
Labridae Doratonotus megalepis ott763958 P 109 1.3 1.2 1.42 No No Yes
Labridae Epibulus insidiator ott120301 P 110 1.29 0.92 1.66 No Yes No
Labridae Gomphosus varius ott358606 P 111 1.32 0.94 1.7 No Yes No
Labridae Halichoeres biocellatus ott1015700 P 112 1.27 1.01 1.54 No Yes No
Labridae Halichoeres bivittatus ott694396 P 113 1.3 1.2 1.4 No No Yes
Labridae Halichoeres chierchiae ott615862 P 114 1.27 1.18 1.38 No No Yes
Labridae Halichoeres chloropterus ott1015715 P 115 1.29 1.19 1.41 No No Yes
Labridae Halichoeres dispilus ott762170 P 116 1.27 1.18 1.38 No No Yes
Labridae Halichoeres garnoti ott665840 P 117 1.28 1.18 1.39 No No Yes
Labridae Halichoeres hortulanus ott694390 P 118 1.34 0.88 1.82 No Yes Yes
Labridae Halichoeres maculipinna ott955368 P 119 1.29 1.19 1.41 No No Yes
Labridae Halichoeres margaritaceus ott694405 P 120 1.29 1.2 1.38 No No Yes
Labridae Halichoeres marginatus ott694408 P 121 1.26 1.17 1.36 No No Yes
Labridae Halichoeres melanurus ott627330 P 122 1.3 0.95 1.69 No Yes Yes
Labridae Halichoeres miniatus ott694399 P 123 1.3 1.2 1.41 No No Yes
Labridae Halichoeres nebulosus ott1015709 P 124 1.29 1.19 1.41 No No Yes
28
Table S7 Continued.
Family Species OTL SM F3 b1 2.5% C.I. 97.5% C.I. D1 D2 D3
Labridae Halichoeres nicholsi ott694402 P 125 1.28 1.18 1.39 No No Yes
Labridae Halichoeres notospilus ott694414 P 126 1.32 1.21 1.43 No No Yes
Labridae Halichoeres pictus ott173446 P 127 1.32 1.21 1.45 No No Yes
Labridae Halichoeres poeyi ott38828 P 128 1.28 1.18 1.39 No No Yes
Labridae Halichoeres trimaculatus ott942026 P 129 1.3 1.19 1.42 No No Yes
Labridae Hemigymnus fasciatus ott924674 P 130 1.29 0.93 1.67 No Yes Yes
Labridae Hemigymnus melapterus ott302368 P 131 1.28 1.18 1.39 No No Yes
Labridae Labrichthys unilineatus ott302371 P 132 1.27 0.96 1.58 No Yes No
Labridae Labroides dimidiatus ott302377 P 133 1.27 0.99 1.54 No Yes No
Labridae Labropsis micronesica ott3636703 P 134 1.19 0.92 1.45 No Yes No
Labridae Labropsis xanthonota ott3636702 P 135 1.25 0.99 1.51 No Yes No
Labridae Macropharyngodon meleagris ott552117 P 136 1.26 0.99 1.54 No Yes No
Labridae Novaculichthys taeniourus ott456983 P 137 1.29 0.94 1.68 No Yes Yes
Labridae Oxycheilinus unifasciatus ott352271 P 138 1.29 0.93 1.66 No Yes No
Labridae Pseudocheilinus hexataenia ott712600 P 139 1.29 0.99 1.6 No Yes No
Labridae Stethojulis bandanensis ott201174 P 140 1.29 1.2 1.4 No No Yes
Labridae Symphodus mediterraneus ott730049 D 141 1.29 1.19 1.41 No No Yes
Labridae Symphodus roissali ott876619 D 142 1.21 1.1 1.33 No Yes Yes
Labridae Symphodus rostratus ott553190 D 143 1.29 1.19 1.4 No No Yes
Labridae Symphodus tinca ott787184 D 144 1.31 1.2 1.42 No No Yes
Labridae Thalassoma bifasciatum ott907850 P 145 0.88 0.71 1.05 Yes No Yes
Labridae Thalassoma hardwicke ott102998 P 146 1.28 1.19 1.37 No No Yes
Labridae Thalassoma jansenii ott1094748 P 147 1.29 1.2 1.39 No No Yes
Labridae Thalassoma lucasanum ott1094750 P 148 1.29 1.2 1.39 No No Yes
Labridae Thalassoma lunare ott948286 P 149 1.28 1.19 1.38 No No Yes
Latidae Lates calcarifer ott6362446 P 150 1.15 0.93 1.38 Yes No No
Liparidae Careproctus reinhardti ott1065176 D 285 1.32 0.62 2.03 Yes Yes No
Liparidae Liparis fabricii ott496423 D 286 1.36 1.02 1.71 No Yes No
Liparidae Paraliparis bathybius ott309222 BM 287 1.31 0.64 1.98 Yes Yes No
Lutjanidae Lutjanus campechanus ott642863 P 151 1.1 0.72 1.47 Yes No No
Lutjanidae Lutjanus carponotatus ott333348 P 152 1.84 1.53 2.14 Yes Yes No
Lutjanidae Lutjanus synagris ott943184 P 153 0.77 0.56 0.99 Yes No No
Lutjanidae Ocyurus chrysurus ott943183 P 154 1.36 0.98 1.73 Yes No No
Lutjanidae Rhomboplites aurorubens ott642869 P 155 1.49 1.34 1.65 Yes No No
Malacanthidae Caulolatilus microps ott39883 P 156 1.63 1.19 2.07 Yes Yes No
Merlucciidae Merluccius gayi ott323082 P 42 1.3 1 1.59 Yes No No
Merlucciidae Merluccius hubbsi ott757316 P 43 1.17 1.08 1.27 Yes No No
Merlucciidae Merluccius merluccius ott5860466 P 44 1.12 0.94 1.3 Yes Yes No
Monacanthidae Brachaluteres jacksonianus ott333693 S 336 1.19 0.76 1.6 Yes Yes No
Monacanthidae Eubalichthys bucephalus ott3631310 S 337 1.42 0.73 2.14 Yes Yes No
Monacanthidae Oxymonacanthus longirostris ott333706 S 338 1.32 0.85 1.8 Yes Yes No
Monacanthidae Paramonacanthus japonicus ott3631319 S 339 1.28 1.02 1.54 No Yes No
Monacanthidae Rudarius ercodes ott511593 S 340 0.92 0.51 1.34 Yes Yes No
Moronidae Dicentrarchus labrax ott3549 P 157 1.34 0.96 1.72 Yes No Yes
29
Table S7 Continued.
Family Species OTL SM F3 b1 2.5% C.I. 97.5% C.I. D1 D2 D3
Mugilidae Chelon labrosus ott777079 P 46 1.06 0.46 1.65 Yes Yes No
Mugilidae Chelon ramada ott668241 P 47 1.24 0.59 1.9 Yes Yes No
Mugilidae Mugil cephalus ott418648 P 48 0.93 0.61 1.23 Yes Yes No
Mugilidae Mugil curema ott418643 P 49 1.26 0.71 1.8 Yes No No
Mugilidae Mugil hospes ott779937 P 50 1.25 0.69 1.81 Yes No No
Mugilidae Mugil liza ott779933 P 51 1.3 0.72 1.88 Yes No No
Mugilidae Mugil trichodon ott246901 P 52 1.29 0.73 1.86 Yes No No
Mugilidae Planiliza subviridis ott823526 P 53 1.1 0.61 1.59 Yes Yes No
Mullidae Parupeneus multifasciatus ott295485 P 158 1.29 1.19 1.41 No No Yes
Muraenesocidae Muraenesox cinereus ott479856 P 1 1.27 1.18 1.35 No No Yes
Nemipteridae Scolopsis bilineata ott463069 P 159 1.29 1.19 1.41 No No Yes
Nototheniidae Lepidonotothen nudifrons ott260052 D 160 1.38 0.69 2.04 Yes Yes No
Nototheniidae Patagonotothen longipes ott430554 D 161 1.44 1.15 1.74 No Yes No
Nototheniidae Patagonotothen sima ott874773 D 162 1.53 1.3 1.78 No Yes No
Osmeridae Mallotus villosus ott225330 D 54 1.33 0.81 1.85 Yes No No
Paralichthyidae Paralichthys dentatus ott1019735 P 259 1.18 0.83 1.52 Yes Yes No
Paralichthyidae Paralichthys patagonicus ott654479 P 260 1.49 1.02 1.97 Yes No No
Pinguipedidae Parapercis snyderi ott3635246 P 163 1.28 1.18 1.38 No No Yes
Plesiopidae Acanthoclinus fuscus ott3633802 D 164 1.57 1.04 2.12 Yes No No
Pleuronectidae Ammotretis rostratus ott600529 P 261 1.27 0.93 1.62 No Yes No
Pleuronectidae Hippoglossoides platessoides ott223665 P 262 1.07 0.93 1.21 Yes No No
Pleuronectidae Hippoglossus hippoglossus ott408328 P 263 1.3 1.21 1.39 No No Yes
Pleuronectidae Pleuronectes platessa ott408337 P 264 1.21 1.12 1.3 Yes No Yes
Pleuronectidae Pseudopleuronectes americanus* ott261419 D 265 1.03 0.65 1.41 Yes Yes Yes
Pleuronectidae Reinhardtius hippoglossoides ott835141 P 266 1.41 1.16 1.65 Yes No No
Pleuronectidae Rhombosolea tapirina ott867269 P 267 1.27 0.95 1.59 No Yes No
Pomacanthidae Pomacanthus zonipectus ott826472 P 165 1.36 0.78 1.93 Yes No No
Pomacentridae Abudefduf bengalensis ott318931 D 166 1.31 1.21 1.43 No No Yes
Pomacentridae Abudefduf saxatilis* ott405751 D 167 0.96 0.39 1.52 Yes Yes Yes
Pomacentridae Abudefduf septemfasciatus ott129790 D 168 1.29 1.19 1.4 No No Yes
Pomacentridae Abudefduf sordidus ott1053071 D 169 1.29 1.2 1.41 No No Yes
Pomacentridae Abudefduf troschelii ott961357 D 170 1.31 1.21 1.42 No No Yes
Pomacentridae Abudefduf vaigiensis ott1053067 D 171 1.29 1.19 1.4 No No Yes
Pomacentridae Acanthochromis polyacanthus ott100410 D 172 1.52 0.95 2.09 Yes No No
Pomacentridae Chromis atrilobata ott436999 D 173 1.34 1.23 1.45 No No Yes
Pomacentridae Chromis atripectoralis ott741423 D 174 1.28 1.19 1.37 No No Yes
Pomacentridae Chromis multilineata ott437016 D 175 1.3 1.21 1.4 No No Yes
Pomacentridae Hypsypops rubicundus ott847666 D 176 1.3 1.21 1.4 No No Yes
Pomacentridae Microspathodon bairdii ott237630 D 177 1.29 1.19 1.39 No No Yes
Pomacentridae Microspathodon chrysurus ott847660 D 178 1.29 1.19 1.39 No No Yes
Pomacentridae Microspathodon dorsalis ott205758 D 179 1.29 1.2 1.39 No No Yes
Pomacentridae Pomacentrus coelestis ott622060 D 180 1.08 0.73 1.43 Yes Yes No
Pomacentridae Stegastes acapulcoensis ott3635554 D 181 1.32 1.22 1.43 No No Yes
Pomacentridae Stegastes adustus ott323181 D 182 1.32 1.22 1.43 No No Yes
30
Table S7 Continued.
Family Species OTL SM F3 b1 2.5% C.I. 97.5% C.I. D1 D2 D3
Pomacentridae Stegastes diencaeus ott729161 D 183 1.28 1.19 1.39 No No Yes
Pomacentridae Stegastes flavilatus ott3635541 D 184 1.31 1.21 1.41 No No Yes
Pomacentridae Stegastes fuscus ott3635543 D 185 1.07 0.73 1.41 Yes No No
Pomacentridae Stegastes leucostictus ott100830 D 186 1.31 1.2 1.43 No No Yes
Pomacentridae Stegastes partitus ott345269 D 187 1.32 1.22 1.42 No No Yes
Pomacentridae Stegastes planifrons ott665837 D 188 1.3 1.2 1.41 No No Yes
Pomacentridae Stegastes variabilis ott323173 D 189 1.32 1.21 1.44 No No Yes
Pristigasteridae Ilisha africana ott741177 P 28 1.4 0.79 2.01 Yes Yes No
Scaridae Cetoscarus bicolor ott1041720 P 190 1.29 0.95 1.63 No Yes No
Scaridae Chlorurus sordidus ott436737 P 191 1.28 0.95 1.65 No Yes Yes
Scaridae Cryptotomus roseus ott321483 P 192 1.32 1.21 1.44 No No Yes
Scaridae Hipposcarus longiceps ott465039 P 193 1.3 0.94 1.67 No Yes No
Scaridae Scarus flavipectoralis ott1041716 P 194 1.28 1.19 1.39 No No Yes
Scaridae Scarus forsteni ott548869 P 195 1.29 0.93 1.66 No Yes No
Scaridae Scarus frenatus ott436725 P 196 1.27 0.93 1.63 No Yes Yes
Scaridae Scarus globiceps ott514869 P 197 1.27 0.88 1.68 No Yes Yes
Scaridae Scarus iseri ott686634 P 198 1.31 1.2 1.43 No No Yes
Scaridae Scarus niger ott14311 P 199 1.26 1.17 1.36 No No Yes
Scaridae Scarus psittacus ott514872 P 200 1.33 1.22 1.46 No No Yes
Scaridae Scarus schlegeli ott514881 P 201 1.26 0.95 1.61 No Yes Yes
Scaridae Scarus spinus ott419732 P 202 1.29 1.19 1.4 No No Yes
Scaridae Sparisoma atomarium ott686627 P 203 1.33 1.22 1.46 No No Yes
Scaridae Sparisoma aurofrenatum ott686624 P 204 1.26 1.17 1.37 No No Yes
Scaridae Sparisoma radians ott214109 P 205 1.29 1.2 1.39 No No Yes
Scaridae Sparisoma rubripinne ott686642 P 206 1.26 1.17 1.37 No No Yes
Sciaenidae Bairdiella chrysoura ott761907 P 207 1.13 0.5 1.75 Yes Yes No
Sciaenidae Cynoscion regalis* ott5860595 P 208 1.51 1.23 1.79 Yes Yes Yes
Sciaenidae Cynoscion striatus ott267367 P 209 1.33 0.77 1.91 Yes No No
Sciaenidae Genyonemus lineatus ott1027240 P 210 2.02 1.76 2.28 Yes No No
Sciaenidae Isopisthus parvipinnis ott133138 P 211 1.42 0.98 1.87 Yes No No
Sciaenidae Larimus breviceps ott924070 P 212 1.82 1.3 2.36 Yes No No
Sciaenidae Larimus fasciatus ott897432 P 213 1.17 0.83 1.51 Yes Yes No
Sciaenidae Leiostomus xanthurus ott1052160 P 214 1.27 1.17 1.38 No No Yes
Sciaenidae Macrodon ancylodon ott99076 P 215 1.35 0.99 1.7 Yes No No
Sciaenidae Micropogonias furnieri ott742890 P 216 1.1 0.92 1.28 Yes No No
Sciaenidae Micropogonias undulatus ott987393 P 217 1.28 1.18 1.4 No No Yes
Sciaenidae Paralonchurus brasiliensis ott160812 P 218 1.01 0.53 1.47 Yes No No
Sciaenidae Seriphus politus ott827210 P 219 1.22 1.09 1.34 Yes No No
Sciaenidae Stellifer rastrifer ott362452 P 220 0.87 0.61 1.15 Yes No No
Scombridae Auxis rochei ott713972 P 221 1.28 0.58 1.98 Yes Yes No
Scombridae Euthynnus affinis ott308760 P 222 1.22 0.9 1.52 No Yes No
Scombridae Katsuwonus pelamis ott308754 P 223 1.36 1.09 1.63 No Yes No
Scombridae Sarda sarda ott833175 P 224 1.35 0.99 1.73 No Yes No
Scombridae Scomber scombrus ott70561 P 225 1.49 1.18 1.8 Yes No No
31
Table S7 Continued.
Family Species OTL SM F3 b1 2.5% C.I. 97.5% C.I. D1 D2 D3
Scombridae Scomberomorus cavalla ott70546 P 226 1.48 1.15 1.82 Yes Yes No
Scombridae Scomberomorus maculatus ott266387 P 227 1.23 0.94 1.52 Yes No No
Scombridae Thunnus alalunga ott833194 P 228 1.04 0.65 1.42 Yes No No
Scombridae Thunnus albacares ott833188 P 229 1.05 0.61 1.5 Yes Yes No
Scombridae Thunnus obesus ott358613 P 230 1.31 1.06 1.57 No Yes No
Scombridae Thunnus thynnus ott358607 P 231 1.17 0.54 1.77 Yes Yes No
Sebastidae Sebastes alutus ott480735 BI 288 1.87 1.7 2.05 Yes No No
Sebastidae Sebastes atrovirens ott480732 BI 289 1.22 0.69 1.73 Yes No No
Sebastidae Sebastes auriculatus ott480729 BI 290 1.17 0.84 1.51 Yes No No
Sebastidae Sebastes brevispinis ott1057081 BI 291 1.39 0.92 1.86 Yes No No
Sebastidae Sebastes carnatus ott1057074 BI 292 1.29 0.76 1.82 Yes No No
Sebastidae Sebastes caurinus ott1057078 BI 293 1.78 1.59 1.99 Yes No No
Sebastidae Sebastes chlorostictus ott1057068 BI 294 1.66 1.44 1.88 Yes No No
Sebastidae Sebastes constellatus ott1057062 BI 295 1.44 1.05 1.84 Yes No No
Sebastidae Sebastes crameri ott1057065 BI 296 1.57 1.2 1.94 Yes No No
Sebastidae Sebastes dallii ott1088425 BI 297 1.25 0.84 1.67 Yes No No
Sebastidae Sebastes diploproa ott1057059 BI 298 1.56 1.24 1.88 Yes No No
Sebastidae Sebastes elongatus ott531065 BI 299 1.4 1.16 1.65 Yes No No
Sebastidae Sebastes entomelas ott531068 BI 300 1.73 1.53 1.93 Yes No No
Sebastidae Sebastes flavidus ott531059 BI 301 1.61 1.35 1.86 Yes No No
Sebastidae Sebastes goodei ott531047 BI 302 1.42 1.23 1.62 Yes No No
Sebastidae Sebastes helvomaculatus ott531044 BI 303 1.4 0.91 1.92 Yes No No
Sebastidae Sebastes hopkinsi ott9589 BI 304 1.12 0.76 1.48 Yes No No
Sebastidae Sebastes jordani ott9592 BI 305 1.21 0.85 1.57 Yes No No
Sebastidae Sebastes levis ott9598 BI 306 1.17 0.89 1.44 Yes No No
Sebastidae Sebastes melanops ott9604 BI 307 1.41 1.15 1.66 Yes No No
Sebastidae Sebastes melanostomus ott9607 BI 308 1.52 1.21 1.84 Yes No No
Sebastidae Sebastes mentella ott1088432 BI 309 1.56 1.17 1.95 Yes Yes No
Sebastidae Sebastes miniatus ott9611 BI 310 1.69 1.47 1.9 Yes No No
Sebastidae Sebastes mystinus ott9616 BI 311 1.85 1.67 2.01 Yes No No
Sebastidae Sebastes norvegicus ott1088441 BI 312 1.16 0.75 1.56 Yes Yes No
Sebastidae Sebastes ovalis ott582324 BI 313 1.35 0.93 1.75 Yes No No
Sebastidae Sebastes paucispinis ott582321 BI 314 1.46 1.28 1.65 Yes No No
Sebastidae Sebastes pinniger ott582318 BI 315 1.36 0.9 1.82 Yes No No
Sebastidae Sebastes rastrelliger ott582315 BI 316 1.34 0.92 1.76 Yes No No
Sebastidae Sebastes rosaceus ott212767 BI 317 1.25 0.94 1.56 Yes No No
Sebastidae Sebastes rosenblatti ott582311 BI 318 1.4 1.08 1.73 Yes No No
Sebastidae Sebastes rufus ott884513 BI 319 1.41 1.03 1.78 Yes No No
Sebastidae Sebastes saxicola ott884510 BI 320 1.15 1.03 1.26 Yes No No
Sebastidae Sebastes semicinctus ott795707 BI 321 1.4 1.09 1.72 Yes No No
Sebastidae Sebastes serranoides ott884519 BI 322 1.66 1.43 1.89 Yes No No
Sebastidae Sebastes viviparus ott99016 BI 323 1.32 1.03 1.6 Yes No No
Sebastidae Sebastolobus altivelis ott811927 P 324 1.28 0.91 1.66 No Yes No
32
Table S7 Continued.
Family Species OTL SM F3 b1 2.5% C.I. 97.5% C.I. D1 D2 D3
Serranidae Cephalopholis cruentata ott12880 P 232 1.29 0.68 1.89 Yes Yes No
Serranidae Epinephelus aeneus ott306171 P 233 1.25 0.97 1.54 Yes No No
Serranidae Epinephelus fasciatus ott534146 P 234 0.9 0.45 1.34 Yes No Yes
Serranidae Epinephelus marginatus ott733897 P 235 1.32 0.76 1.89 Yes No No
Serranidae Epinephelus tauvina ott218649 P 236 0.94 0.41 1.45 Yes Yes No
Serranidae Mycteroperca tigris ott207413 P 237 1.54 1.05 2.05 Yes No No
Serranidae Serranus tabacarius ott284266 P 238 1.29 1.2 1.41 No No Yes
Serranidae Serranus tigrinus ott807897 P 239 1.26 1.16 1.36 No No Yes
Serranidae Serranus tortugarum ott511782 P 240 1.29 1.19 1.41 No No Yes
Siganidae Siganus canaliculatus ott199059 S 241 1.4 0.98 1.82 Yes Yes No
Soleidae Solea solea* ott710233 P 268 1.26 1.02 1.52 Yes Yes Yes
Sparidae Acanthopagrus latus ott760733 P 242 1.26 0.57 1.94 Yes Yes No
Sparidae Archosargus rhomboidalis ott724892 P 243 1.14 0.74 1.54 Yes Yes No
Sparidae Sparidentex hasta ott845300 P 244 1.33 0.74 1.91 Yes Yes No
Stromateidae Pampus chinensis ott916290 P 245 1.24 0.55 1.93 Yes Yes No
Syngnathidae Entelurus aequoreus ott971459 BP 325 1.4 1.19 1.61 No Yes No
Syngnathidae Hippocampus reidi ott630138 BP 326 1.35 0.88 1.83 Yes No No
Syngnathidae Micrognathus crinitus ott3642321 BP 327 1.21 0.86 1.57 Yes Yes No
Syngnathidae Nerophis ophidion ott806821 BP 328 1.31 1.13 1.5 No Yes No
Syngnathidae Syngnathus acus ott774440 BP 329 1.34 1.18 1.51 No Yes No
Syngnathidae Syngnathus floridae ott774435 BP 330 1.44 0.95 1.92 Yes Yes No
Syngnathidae Syngnathus louisianae ott774414 BP 331 1.48 0.95 2 Yes Yes No
Syngnathidae Syngnathus rostellatus ott774411 BP 332 1.42 1.26 1.58 No Yes No
Syngnathidae Syngnathus scovelli ott251400 BP 333 1.37 0.95 1.79 Yes Yes No
Syngnathidae Syngnathus typhle ott251396 BP 334 1.32 1.19 1.45 No Yes No
Tetraodontidae Canthigaster rostrata ott484093 D 341 1.29 1.19 1.4 No No Yes
Tetraodontidae Canthigaster valentini ott701127 S 342 0.98 0.65 1.31 Yes Yes No
Trachichthyidae Hoplostethus atlanticus ott816600 P 9 1.12 0.91 1.33 Yes No No
Trichiuridae Aphanopus carbo ott113465 P 246 1.32 0.74 1.91 Yes Yes No
Trichodontidae Arctoscopus japonicus ott883978 D 247 1.29 1.2 1.39 No No Yes
Zoarcidae Gymnelus viridis ott729337 D 248 1.34 0.8 1.88 Yes Yes No
Zoarcidae Lycodes esmarkii ott841648 D 249 1.04 0.41 1.68 Yes Yes No
Zoarcidae Lycodes eudipleurostictus ott905435 D 250 1.33 0.64 2.01 Yes Yes No
Zoarcidae Lycodes frigidus ott539690 D 251 1.23 0.52 1.93 Yes Yes No
Zoarcidae Lycodes pallidus ott178074 D 252 1.43 0.82 2.05 Yes Yes No
Zoarcidae Lycodes polaris ott34255 D 253 1.31 0.97 1.64 No Yes No
Zoarcidae Lycodes reticulatus ott752411 D 254 1.21 0.83 1.61 No Yes No
Zoarcidae Lycodes vahlii ott178063 D 255 1.16 0.66 1.66 Yes Yes No
Zoarcidae Melanostigma atlanticum ott3633033 D 256 1.1 0.63 1.58 Yes Yes No
Zoarcidae Pachycara brachycephalum ott203179 D 257 1.27 0.9 1.63 No Yes No
The above results were obtained by combining the posterior distributions of parameter estimates from Models 1 (Table S3), 2 (Table S4), and 3 (Table S5) as described in the Materials and Methods Section. If a species was present in given
dataset (i.e. fecundity, egg-volume, and/or egg-energy), we used the species-specific posterior distributions of parameter estimates as obtained using random effects. However, if the species was not present, we used the overall fixed-effects
posterior distributions. Columns D1–D3 above indicate in which dataset a species was present. Only 9 species were present in all three datasets (marked with an *): Abudefduf saxatilis, Cynoscion regalis, Engraulis mordax, Gadus morhua,
Gasterosteus aculeatus, Ostorhinchus cyanosoma, Oxylebius pictus, Pseudopleuronectes americanus, Solea solea. This is the same species list used to draw Fig. 3 in the main text.
33
Table S8. Summary statistics for ordinary least squares regressions between total clutch
dry weight (g) and female mass (g) for three studies that measured egg dry weight, fecundity
and female size. The equation is: Clutch dry weight = b0  Female massb1 . Model fitting was
conducted by taking the natural logarithm from both sides of the equation (i.e. equivalent to
equation 1 in Materials and Methods). b0 = intercept; b1 = Mean scaling exponent; b1 S.E. =
Standard error of mean scaling exponent; n = number of independent measurements for each
species.
Species b0 b1 b1 S.E. n Refs
Pseudopleuronectes
americanus 0.01 1.32 0.12 17 (152)
Clupea harengus 0.02 1.01 0.11 45 (243)
Pomacentrus coelestis 0 1.54 0.09 247 (157)
34
References and Notes 
1. R. E. Earll, in National Oceanic and Atmospheric Administration Report of the United States 
Commissioner of Fisheries, S. F. Baird, Ed. (Government printing office, San Diego, 
1881), pp. 395–426. 
2. R. J. H. Beverton, S. J. Holt, On the Dynamics of Exploited Fish Populations (Fishery 
Investigations Series II, Her Majesty’s Stationery Office, London, 1957), vol. 19. 
3. C. T. Marshall, C. L. Needle, A. Thorsen, O. S. Kjesbu, N. A. Yaragina, Systematic bias in 
estimates of reproductive potential of an Atlantic cod (Gadus morhua) stock: 
Implications for stock–recruit theory and management. Can. J. Fish. Aquat. Sci. 63, 980–
994 (2006). doi:10.1139/f05-270 
4. M. A. Hixon, D. W. Johnson, S. M. Sogard, BOFFFFs: On the importance of conserving old-
growth age structure in fishery populations. ICES J. Mar. Sci. 71, 2171–2185 (2014). 
doi:10.1093/icesjms/fst200 
5. J. N. Lim, A. M. Senior, S. Nakagawa, Heterogeneity in individual quality and reproductive 
trade-offs within species. Evolution 68, 2306–2318 (2014). doi:10.1111/evo.12446 
Medline 
6. C. Birkeland, P. K. Dayton, The importance in fishery management of leaving the big ones. 
Trends Ecol. Evol. 20, 356–358 (2005). doi:10.1016/j.tree.2005.03.015 Medline 
7. S. E. Lester, B. S. Halpern, K. Grorud-Colvert, J. Lubchenco, B. I. Ruttenberg, S. D. Gaines, 
S. Airamé, R. R. Warner, Biological effects within no-take marine reserves: A global 
synthesis. Mar. Ecol. Prog. Ser. 384, 33–46 (2009). doi:10.3354/meps08029 
8. Materials and methods are available as supplementary materials. 
9. R. Beldade, S. J. Holbrook, R. J. Schmitt, S. Planes, D. Malone, G. Bernardi, Larger female 
fish contribute disproportionately more to self-replenishment. Proc. R. Soc. Lond. B Biol. 
Sci. 279, 2116–2121 (2012). doi:10.1098/rspb.2011.2433 Medline 
10. I. van Rijn, Y. Buba, J. DeLong, M. Kiflawi, J. Belmaker, Large but uneven reduction in fish 
size across species in relation to changing sea temperatures. Glob. Chang. Biol. 23, 3667–
3674 (2017). doi:10.1111/gcb.13688 Medline 
11. D. R. Barneche, M. Kulbicki, S. R. Floeter, A. M. Friedlander, J. Maina, A. P. Allen, Scaling 
metabolism from individuals to reef-fish communities at broad spatial scales. Ecol. Lett. 
17, 1067–1076 (2014). doi:10.1111/ele.12309 Medline 
12. K. H. Andersen, J. L. Blanchard, E. A. Fulton, H. Gislason, N. S. Jacobsen, T. van Kooten, 
Assumptions behind size-based ecosystem models are realistic. ICES J. Mar. Sci. 73, 
1651–1655 (2016). doi:10.1093/icesjms/fsv211 
13. R. Froese, C. Walters, D. Pauly, H. Winker, O. L. F. Weyl, N. Demirel, A. C. Tsikliras, S. J. 
Holt, A critique of the balanced harvesting approach to fishing. ICES J. Mar. Sci. 73, 
1640–1650 (2016). doi:10.1093/icesjms/fsv122 
14. M. Heino, L. Baulier, D. S. Boukal, B. Ernande, F. D. Johnston, F. M. Mollet, H. Pardoe, N. 
O. Therkildsen, S. Uusi-Heikkila, A. Vainikka, R. Arlinghaus, D. J. Dankel, E. S. 
Dunlop, A. M. Eikeset, K. Enberg, G. H. Engelhard, C. Jorgensen, A. T. Laugen, S. 
35
Matsumura, S. Nussle, D. Urbach, R. Whitlock, A. D. Rijnsdorp, U. Dieckmann, Can 
fisheries-induced evolution shift reference points for fisheries management? ICES J. Mar. 
Sci. 70, 707–721 (2013). doi:10.1093/icesjms/fst077 
15. D. J. Marshall, S. S. Heppell, S. B. Munch, R. R. Warner, The relationship between maternal 
phenotype and offspring quality: Do older mothers really produce the best offspring? 
Ecology 91, 2862–2873 (2010). doi:10.1890/09-0156.1 Medline 
16. C. J. Harvey, N. Tolimieri, P. S. Levin, Changes in body size, abundance, and energy 
allocation in rockfish assemblages of the northeast Pacific. Ecol. Appl. 16, 1502–1515 
(2006). doi:10.1890/1051-0761(2006)016[1502:CIBSAA]2.0.CO;2 Medline 
17. M. J. Genner, D. W. Sims, A. J. Southward, G. C. Budd, P. Masterson, M. McHugh, P. 
Rendle, E. J. Southall, V. J. Wearmouth, S. J. Hawkins, Body size-dependent responses 
of a marine fish assemblage to climate change and fishing over a century-long scale. 
Glob. Change Biol. 16, 517–527 (2010). doi:10.1111/j.1365-2486.2009.02027.x 
18. R. A. Fritzsche, Development of Fishes of the Mid-Atlantic Bight: An Atlas of Egg, Larval, 
and Juvenile Stages (Fish and Wildlife Service, U.S. Department of the Interior, 1978), 
vol. 5. 
19. C. M. Duarte, M. Alcaraz, To produce many small or few large eggs: A size-independent 
reproductive tactic of fish. Oecologia 80, 401–404 (1989). doi:10.1007/BF00379043 
Medline 
20. M. A. Elgar, Evolutionary compromise between a few large and many small eggs: 
Comparative evidence in teleost fish. Oikos 59, 283–287 (1990). doi:10.2307/3545546 
21. H. G. Moser, The Early Stages of Fishes in the California Current Region. Atlas No. 33 
(Allen Press, Lawrence, KS, 1996). 
22. S. Einum, I. A. Fleming, Does within-population variation in fish egg size reflect maternal 
influences on optimal values? Am. Nat. 160, 756–765 (2002). Medline 
23. K. Kasimatis, C. Riginos, A phylogenetic analysis of egg size, clutch size, spawning mode, 
adult body size, and latitude in reef fishes. Coral Reefs 35, 387–397 (2016). 
doi:10.1007/s00338-015-1380-1 
24. R. Froese, D. Pauly, FishBase (2017); www.fishbase.org. 
25. R. Froese, J. T. Thorson, R. B. Reyes Jr., A Bayesian approach for estimating length-weight 
relationships in fishes. J. Appl. Ichthyology 30, 78–85 (2014). doi:10.1111/jai.12299 
26. D. R. Robertson, R. Collin, Inter- and intra-specific variation in egg size among reef fishes 
across the Isthmus of Panama. Front. Ecol. Evol. 2, 84 (2015). 
doi:10.3389/fevo.2014.00084 
27. E. Kamler, Parent–egg–progeny relationships in teleost fishes: An energetics perspective. 
Rev. Fish Biol. Fish. 15, 399–421 (2005). doi:10.1007/s11160-006-0002-y 
28. M. Lynch, Methods for the analysis of comparative data in evolutionary biology. Evolution 
45, 1065–1080 (1991). doi:10.1111/j.1558-5646.1991.tb04375.x Medline 
29. M. Pagel, Inferring the historical patterns of biological evolution. Nature 401, 877–884 
(1999). doi:10.1038/44766 Medline 
36
30. J. D. Hadfield, S. Nakagawa, General quantitative genetic methods for comparative biology: 
Phylogenies, taxonomies and multi-trait models for continuous and categorical 
characters. J. Evol. Biol. 23, 494–508 (2010). doi:10.1111/j.1420-9101.2009.01915.x 
Medline 
31. F. Michonneau, J. W. Brown, D. J. Winter, rotl: An R package to interact with the Open Tree 
of Life data. Methods Ecol. Evol. 7, 1476–1481 (2016). doi:10.1111/2041-210X.12593 
32. E. A. Housworth, E. P. Martins, M. Lynch, The phylogenetic mixed model. Am. Nat. 163, 
84–96 (2004). doi:10.1086/380570 Medline 
33. B. Frédérich, L. Sorenson, F. Santini, G. J. Slater, M. E. Alfaro, Iterative ecological radiation 
and convergence during the evolutionary history of damselfishes (Pomacentridae). Am. 
Nat. 181, 94–113 (2013). doi:10.1086/668599 Medline 
34. R. Betancur-R, R. E. Broughton, E. O. Wiley, K. Carpenter, J. A. López, C. Li, N. I. 
Holcroft, D. Arcila, M. Sanciangco, J. C. Cureton Ii, F. Zhang, T. Buser, M. A. 
Campbell, J. A. Ballesteros, A. Roa-Varon, S. Willis, W. C. Borden, T. Rowley, P. C. 
Reneau, D. J. Hough, G. Lu, T. Grande, G. Arratia, G. Ortí, The tree of life and a new 
classification of bony fishes. PLOS Curr. 5, 1 (2013). 
doi:10.1371/currents.tol.53ba26640df0ccaee75bb165c8c26288 Medline 
35. A. Grafen, The phylogenetic regression. Philos. Trans. R. Soc. Lond. B Biol. Sci. 326, 119–
157 (1989). doi:10.1098/rstb.1989.0106 Medline 
36. E. Paradis, J. Claude, K. Strimmer, APE: Analyses of phylogenetics and evolution in R 
language. Bioinformatics 20, 289–290 (2004). doi:10.1093/bioinformatics/btg412 
Medline 
37. P. Bürkner, brms: An R package for Bayesian multilevel models using Stan. J. Stat. Softw. 
80, 1–28 (2017). 
38. M. B. Hooten, N. T. Hobbs, A guide to Bayesian model selection for ecologists. Ecol. 
Monogr. 85, 3–28 (2015). doi:10.1890/14-0661.1 
39. A. Vehtari, A. Gelman, J. Gabry, loo: Efficient leave-one-out cross-validation and WAIC for 
Bayesian models (2016); https://github.com/jgabry/loo. 
40. M. Gadgil, W. H. Bossert, Life historical consequences of natural selection. Am. Nat. 104, 1–
24 (1970). doi:10.1086/282637 
41. D. A. Roff, An allocation model of growth and reproduction in fish. Can. J. Fish. Aquat. Sci. 
40, 1395–1404 (1983). doi:10.1139/f83-161 
42. D. A. Roff, The evolution of life history parameters in teleosts. Can. J. Fish. Aquat. Sci. 41, 
989–1000 (1984). doi:10.1139/f84-114 
43. M. J. Reiss, The allometry of reproduction: Why larger species invest relatively less in their 
offspring. J. Theor. Biol. 113, 529–544 (1985). doi:10.1016/S0022-5193(85)80037-0 
44. J. Kozłowski, J. Uchmanski, Optimal individual growth and reproduction in perennial 
species with indeterminate growth. Evol. Ecol. 1, 214–230 (1987). 
doi:10.1007/BF02067552 
37
45. J. Kozłowski, Optimal allocation of resources explains interspecific life-history patterns in 
animals with indeterminate growth. Proc. R. Soc. London Ser. B 263, 559–566 (1996). 
doi:10.1098/rspb.1996.0084 
46. G. B. West, J. H. Brown, B. J. Enquist, A general model for ontogenetic growth. Nature 413, 
628–631 (2001). doi:10.1038/35098076 Medline 
47. E. L. Charnov, T. F. Turner, K. O. Winemiller, Reproductive constraints and the evolution of 
life histories with indeterminate growth. Proc. Natl. Acad. Sci. U.S.A. 98, 9460–9464 
(2001). doi:10.1073/pnas.161294498 Medline 
48. E. L. Charnov, Reproductive effort, offspring size and benefit/cost ratios in the classification 
of life histories. Evol. Ecol. Res. 4, 749–758 (2002). 
49. N. P. Lester, B. J. Shuter, P. A. Abrams, Interpreting the von Bertalanffy model of somatic 
growth in fishes: The cost of reproduction. Proc. R. Soc. Lond. B Biol. Sci. 271, 1625–
1631 (2004). doi:10.1098/rspb.2004.2778 Medline 
50. D. A. Roff, E. Heibo, L. A. Vøllestad, The importance of growth and mortality costs in the 
evolution of the optimal life history. J. Evol. Biol. 19, 1920–1930 (2006). 
doi:10.1111/j.1420-9101.2006.01155.x Medline 
51. C. Quince, P. A. Abrams, B. J. Shuter, N. P. Lester, Biphasic growth in fish I: Theoretical 
foundations. J. Theor. Biol. 254, 197–206 (2008). doi:10.1016/j.jtbi.2008.05.029 Medline 
52. L. Pecquerie, P. Petitgas, S. A. L. M. Kooijman, Modeling fish growth and reproduction in 
the context of the dynamic energy budget theory to predict environmental impact on 
anchovy spawning duration. J. Sea Res. 62, 93–105 (2009). 
doi:10.1016/j.seares.2009.06.002 
53. S. A. L. M. Kooijman, Dynamic Energy Budget Theory for Metabolic Organisation 
(Cambridge Univ. Press, 2010). 
54. J. D. Arendt, Size-fecundity relationships, growth trajectories, and the temperature-size rule 
for ectotherms. Evolution 65, 43–51 (2011). doi:10.1111/j.1558-5646.2010.01112.x 
Medline 
55. S. Ohnishi, T. Yamakawa, H. Okamura, T. Akamine, A note on the von Bertalanffy growth 
function concerning the allocation of surplus energy to reproduction. Fish Bull. 110, 223–
229 (2012). 
56. T. Brunel, B. Ernande, F. M. Mollet, A. D. Rijnsdorp, Estimating age at maturation and 
energy-based life-history traits from individual growth trajectories with nonlinear mixed-
effects models. Oecologia 172, 631–643 (2013). doi:10.1007/s00442-012-2527-1 
Medline 
57. E. L. Charnov, H. Gislason, J. G. Pope, Evolutionary assembly rules for fish life histories. 
Fish Fish. 14, 213–224 (2013). doi:10.1111/j.1467-2979.2012.00467.x 
58. D. S. Boukal, U. Dieckmann, K. Enberg, M. Heino, C. Jørgensen, Life-history implications 
of the allometric scaling of growth. J. Theor. Biol. 359, 199–207 (2014). 
doi:10.1016/j.jtbi.2014.05.022 Medline 
38
59. S. A. L. M. Kooijman, K. Lika, Comparative energetics of the 5 fish classes on the basis of 
dynamic energy budgets. J. Sea Res. 94, 19–28 (2014). doi:10.1016/j.seares.2014.01.015 
60. C. V. Minte-Vera, M. N. Maunder, J. M. Casselman, S. E. Campana, Growth functions that 
incorporate the cost of reproduction. Fish. Res. 180, 31–44 (2016). 
doi:10.1016/j.fishres.2015.10.023 
61. M. Jusup, T. Sousa, T. Domingos, V. Labinac, N. Marn, Z. Wang, T. Klanjšček, Physics of 
metabolic organization. Phys. Life Rev. 20, 1–39 (2017). doi:10.1016/j.plrev.2016.09.001 
Medline 
62. M. Mangel, The inverse life-history problem, size-dependent mortality and two extensions of 
results of Holt and Beverton. Fish Fish. 18, 1192–1200 (2017). doi:10.1111/faf.12247 
63. I. M. Smallegange, H. Caswell, M. E. M. Toorians, A. M. de Roos, Mechanistic description 
of population dynamics using dynamic energy budget theory incorporated into integral 
projection models. Methods Ecol. Evol. 8, 146–154 (2017). doi:10.1111/2041-
210X.12675 
64. B. E. Scott, G. Marteinsdottir, G. A. Begg, P. J. Wright, O. S. Kjesbu, Effects of population 
size/age structure, condition and temporal dynamics of spawning on reproductive output 
in Atlantic cod Gadus morhua. Ecol. Modell. 191, 383–415 (2006). 
doi:10.1016/j.ecolmodel.2005.05.015 
65. C. Jørgensen, Ø. Fiksen, State-dependent energy allocation in cod Gadus morhua. Can. J. 
Fish. Aquat. Sci. 63, 186–199 (2006). doi:10.1139/f05-209 
66. K. Enberg, C. Jørgensen, M. Mangel, Fishing-induced evolution and changing reproductive 
ecology of fish: The evolution of steepness. Can. J. Fish. Aquat. Sci. 67, 1708–1719 
(2010). doi:10.1139/F10-090 
67. A. M. Eikeset, A. Richter, E. S. Dunlop, U. Dieckmann, N. C. Stenseth, Economic 
repercussions of fisheries-induced evolution. Proc. Natl. Acad. Sci. U.S.A. 110, 12259–
12264 (2013). doi:10.1073/pnas.1212593110 Medline 
68. N. P. Lester, B. J. Shuter, P. Venturelli, D. Nadeau, Life-history plasticity and sustainable 
exploitation: A theory of growth compensation applied to walleye management. Ecol. 
Appl. 24, 38–54 (2014). doi:10.1890/12-2020.1 Medline 
69. K. H. Andersen, J. E. Beyer, Size structure, not metabolic scaling rules, determines fisheries 
reference points. Fish Fish. 16, 1–22 (2015). doi:10.1111/faf.12042 
70. A. M. Eikeset, E. S. Dunlop, M. Heino, G. Storvik, N. C. Stenseth, U. Dieckmann, Roles of 
density-dependent growth and life history evolution in accounting for fisheries-induced 
trait changes. Proc. Natl. Acad. Sci. U.S.A. 113, 15030–15035 (2016). 
doi:10.1073/pnas.1525749113 Medline 
71. F. Zimmermann, C. Jørgensen, Bioeconomic consequences of fishing-induced evolution: A 
model predicts limited impact on net present value. Can. J. Fish. Aquat. Sci. 72, 612–624 
(2015). doi:10.1139/cjfas-2014-0006 
72. M. Hartvig, K. H. Andersen, J. E. Beyer, Food web framework for size-structured 
populations. J. Theor. Biol. 272, 113–122 (2011). doi:10.1016/j.jtbi.2010.12.006 Medline 
39
73. D. A. Carozza, D. Bianchi, E. D. Galbraith, The ecological module of BOATS-1.0: A 
bioenergetically constrained model of marine upper trophic levels suitable for studies of 
fisheries and ocean biogeochemistry. Geosci. Model Dev. 9, 1545–1565 (2016). 
doi:10.5194/gmd-9-1545-2016 
74. L. Blueweiss, H. Fox, V. Kudzma, D. Nakashima, R. Peters, S. Sams, Relationships between 
body size and some life history parameters. Oecologia 37, 257–272 (1978). 
doi:10.1007/BF00344996 Medline 
75. R. J. Wootton, in Symposia of the Zoological Society of London (Academic Press, 1979), vol. 
44, pp. 133–159. 
76. R. J. Wootton, Constraints in the evolution of fish life histories. Neth. J. Zool. 42, 291–303 
(1992). 
77. A. Hayward, J. F. Gillooly, The cost of sex: Quantifying energetic investment in gamete 
production by males and females. PLOS ONE 6, e16557 (2011). 
doi:10.1371/journal.pone.0016557 Medline 
78. E. J. Dick, S. Beyer, M. Mangel, S. Ralston, A meta-analysis of fecundity in rockfishes 
(genus Sebastes). Fish. Res. 187, 73–85 (2017). doi:10.1016/j.fishres.2016.11.009 
79. T. S. Rass, Geograficheskie parallelizmy v stroenii kostistykh ryb severnykh morei 
(Analogous or parallel variations in structure and development of fishes in Northern and 
Arctic seas) (Moscow Society of Naturalists, MOIP, Moscow, 1941). 
80. L. Fishelson, O. Gon, Comparative oogenesis in cardinal fishes (Apogonidae, Perciformes), 
with special focus on the adaptive structures of the egg envelopes. Environ. Biol. Fishes 
81, 397–414 (2008). doi:10.1007/s10641-007-9211-z 
81. A. W. H. Turnpenny, R. N. Bamber, P. A. Henderson, Biology of the sand-smelt (Atherina 
presbyter Valenciennes) around Fawley power station. J. Fish Biol. 18, 417–427 (1981). 
doi:10.1111/j.1095-8649.1981.tb03783.x 
82. W. F. Thompson, J. B. Thompson, “The spawning of the grunion (Leuresthes tenuis),” in 
Fish Bulletin (California State Printing Office, Sacramento, CA, 1964), pp. 1–30. 
83. A. Moresco, thesis, Fundação Universidade Federal do Rio Grande, Rio Grande, RS, Brazil 
(2006). 
84. H. Kacem, L. Neifar, The reproductive biology of the grey triggerfish Balistes capriscus 
(Pisces: Balistidae) in the Gulf of Gabès (south-eastern Mediterranean Sea). J. Mar. Biol. 
Assoc. U. K. 94, 1531–1537 (2014). doi:10.1017/S0025315414000824 
85. E. T. Lang, G. R. Fitzhugh, Oogenesis and fecundity type of gray triggerfish in the Gulf of 
Mexico. Mar. Coast. Fish. 7, 338–348 (2015). doi:10.1080/19425120.2015.1069428 
86. L. Locatello, F. C. Neat, Reproductive allocation in Aidablennius sphynx (Teleostei, 
Blenniidae): Females lay more eggs faster when paired with larger males. J. Exp. Zool. A 
Comp. Exp. Biol. 303, 922–926 (2005). doi:10.1002/jez.a.204 Medline 
87. R. N. Gibson, I. A. Ezzi, The biology of the scaldfish, Arnoglossus laterna (Walbaum) on the 
west coast of Scotland. J. Fish Biol. 17, 565–575 (1980). doi:10.1111/j.1095-
8649.1980.tb02788.x 
40
88. L. S. J. Hales, Distribution, abundance, reproduction, food habits, age, and growth of round 
scad, Decapterus punctatus, in the South Atlantic Bight. Fish Bull. 85, 251–268 (1987). 
89. P. B. Pinheiro, thesis, Universidade Federal de Pernambuco, Recife, PE, Brazil (2004). 
90. J. de Ciechomski, M. C. Cassia, Reproducción y fecundidad del surel (Trachurus picturatus 
australis). Rev. Invest. Dessaro. Pesq. 2, 39–45 (1980). 
91. V. Caballero-Chávez, Reproducción y fecundidad del robalo blanco (Centropomus 
undecimalis) en el suroeste de Campeche. Cienc. Pesq. 19, 35–45 (2011). 
92. N. Veerappanm, M. Ramanathan, V. Ramaiyan, Maturation and spawning biology of 
Amblygaster sirm from Parangipettai, southeast coast of India. J. Mar. Biol. Assoc. India 
39, 89–96 (1997). 
93. G. J. Óskarsson, C. T. Taggart, Fecundity variation in Icelandic summer-spawning herring 
and implications for reproductive potential. ICES J. Mar. Sci. 63, 493–503 (2006). 
doi:10.1016/j.icesjms.2005.10.002 
94. J. Blay, K. N. Eyeson, Observations on the reproductive biology of the shad, Ethmalosa 
fimbriata (Bowdich), in the coastal waters of Cape Coast, Ghana. J. Fish Biol. 21, 485–
496 (1982). doi:10.1111/j.1095-8649.1982.tb02854.x 
95. C. F. Chubb, I. C. Potter, The reproductive biology and estuarine movements of the gizzard 
shad, Nematalosa vlaminghi (Munro). J. Fish Biol. 25, 527–543 (1984). 
doi:10.1111/j.1095-8649.1984.tb04899.x 
96. J. A. Rodriguez, J. A. Palacios, A. L. Chavarría, Fecundidad y epoca de desove del arenque 
hebra Opisthonema libertate (Pisces: Clupeidae) en el Golfo de Nicoya, Costa Rica. 
Uniciencia 3, 87–93 (1986). 
97. J. A. Rodriguez, J. A. Palacios, A. L. Chavarra, Epoca de maduracin y fecundidad de la 
sardina gallera Opisthonema medirastre y O. bulleri (Pisces: Clupeidae) en la Costa 
Pacfica Central de Costa Rica. Rev. Biol. Trop. 37, 49–54 (1989). 
98. B. L. P. Alvarez, thesis, Centro Interdisciplinario de Ciencias Marinas, Instituto Politecnico 
Nacional, La Paz, Mexico (2008). 
99. P. Dalzell, Some aspects of the reproductive biology of Spratelloides gracilis (Schlegel) in 
the Ysabel Passage, Papua New Guinea. J. Fish Biol. 27, 229–237 (1985). 
doi:10.1111/j.1095-8649.1985.tb04023.x 
100. L. A. Cubillos, C. Alarcon, C. Castillo-Jordan, G. Claramunt, C. Oyarzun, S. Gacitua, 
Spatial and temporal changes in batch fecundity of common sardine and anchovy off 
central and southern Chile (2002–2007). Cienc. Mar. 37 (4B), 547–564 (2011). 
doi:10.7773/cm.v37i4B.1749 
101. E. E. DeMartini, Spatial aspects of reproduction in buffalo sculpin, Enophrys bison. 
Environ. Biol. Fishes 3, 331–336 (1978). doi:10.1007/BF00000524 
102. J. N. Souza, M. T. D. Giamas, H. Vermulm Júnior, Tipo de desova e fecundidade em 
Anchoviella lepidentostole (Fowler, 1911). Revista da Faculdade de Medicina 
Veterinária e Zootecnia da Universidade de São Paulo 25, 251–260 (1988). 
doi:10.11606/issn.2318-3659.v25i2p251-260 
41
103. C. L. Petterson, Fecundity of the anchoveta (Cetengraulis mysticetus) in the Gulf of 
Panama. Bull. I-ATCC VI, 55–68 (1961). 
104. M. Pájaro, G. J. Macchi, R. P. Sánchez, Fecundidad y frecuencia reproductiva de las 
poblaciones bonaerense y patagonica de anchoita argentina (Engraulis anchoite). Rev. 
Invest. Dessaro. Pesq. 11, 19–38 (1997). 
105. J. L. Laroche, S. L. Richardson, Reproduction of northern anchovy, Engraulis mordax, off 
Oregon and Washington. Fish Bull. 7, 603–618 (1980). 
106. A. P. De La Matta, B. B. Díaz, Fecundidad, frecuencia de desove y estado de madurez 
gonadal de la anchovieta peruana durante el invierno austral 2001. Informativo del 
Instituto del Mar del Perú 32, 119–122 (2004). 
107. J. B. Miñano, Estudio de la fecundidad y ciclo sexual de la anchovieta (Engraulis ringens, 
J.) en la zona de Chimbote. Boletín del Instituto del Mar del Perú I, 505–552 (1968). 
108. M. R. Oliveira, A. L. S. Morais, M. M. Carvalho, A. M. Silva, J. T. A. X. Lima, N. T. 
Chellappa, S. Chellappa, Estratégias reprodutivas de sete espécies de peixes das águas 
costeiras do Rio Grande do Norte, Brasil. HOLOS 6, 107–122 (2015). 
doi:10.15628/holos.2015.3601 
109. A. W. May, Fecundity of Atlantic cod. J. Fish. Res. Board Can. 24, 1531–1551 (1967). 
doi:10.1139/f67-127 
110. G. Marteinsdottir, G. A. Begg, Essential relationships incorporating the influence of age, 
size and condition on variables required for estimation of reproductive potential in 
Atlantic cod Gadus morhua. Mar. Ecol. Prog. Ser. 235, 235–256 (2002). 
doi:10.3354/meps235235 
111. O. Kjesbu, P. Witthames, P. Solemdal, M. Greer Walker, Temporal variations in the 
fecundity of Arcto-Norwegian cod (Gadus morhua) in response to natural changes in 
food and temperature. J. Sea Res. 40, 303–321 (1998). doi:10.1016/S1385-
1101(98)00029-X 
112. E. Oosthuizen, N. Daan, Egg fecundity and maturity of North Sea cod, Gadus morhua. 
Neth. J. Sea Res. 8, 378–397 (1974). doi:10.1016/0077-7579(74)90006-4 
113. J. R. G. Hislop, W. B. Hall, The fecundity of whiting, Merlangius merlangus (L.) in the 
North Sea, the Minch and at Iceland. ICES J. Mar. Sci. 36, 42–49 (1974). 
doi:10.1093/icesjms/36.1.42 
114. R. P. Sánchez, J. D. de Ciechomski, E. M. Acha, Estudios sobre reproducción y fecundidad 
de la polaca (Micromesistius australis Norma 1937) en el Mar Argentino. Rev. Invest. 
Dessaro. Pesq. 6, 21–43 (1986). 
115. R. J. Wootton, Fecundity of the three-spined stickleback, Gasterosteus aculeatus (L.). J. 
Fish Biol. 5, 683–688 (1973). doi:10.1111/j.1095-8649.1973.tb04504.x 
116. A. J. Crivelli, R. H. Britton, Life history adaptations of Gasterosteus aculeatus in a 
Mediterranean wetland. Environ. Biol. Fishes 18, 109–125 (1987). 
doi:10.1007/BF00002599 
42
117. R. J. Valenti, The embryology of the neon goby, Gobiosoma oceanops. Copeia 1972, 477–
482 (1972). doi:10.2307/1442919 
118. V. G. Springer, A. J. McErlean, Spawning seasons and growth of the code goby, 
Gobiosoma robustum (Pisces: Gobiidae), in the Tampa Bay area. Tulane Studies in 
Zoology 9, 87–98 (1961). 
119. B. Lassig, Field observations on the reproductive behaviour of Paragobiodon spp. 
(Osteichthyes: Gobiidae) at Heron Island Great Barrier Reef. Mar. Behav. Physiol. 3, 
283–293 (1976). doi:10.1080/10236247609378517 
120. M. C. Healey, Gonad development and fecundity of the sand goby, Gobius minutus Pallas. 
Trans. Am. Fish. Soc. 100, 520–526 (1971). doi:10.1577/1548-
8659(1971)100<520:GDAFOT>2.0.CO;2 
121. H. E. Warfel, D. Merriman, The spawning habits, eggs and larvae of the sea raven, 
Hemitripterus americanus, in southern New England. Copeia 1944, 197–205 (1944). 
doi:10.2307/1438673 
122. E. E. DeMartini, M. E. Anderson, Comparative survivorship and life history of painted 
greenling (Oxylebius pictus) in Puget Sound, Washington and Monterey Bay, California. 
Environ. Biol. Fishes 5, 33–47 (1980). doi:10.1007/BF00000948 
123. T. L. O. Davis, Estimation of fecundity in barramundi, Lates calcarifer (Bloch), using an 
automatic particle counter. Mar. Freshw. Res. 35, 111–118 (1984). 
doi:10.1071/MF9840111 
124. T. C. V. Gesteira, C. T. C. Ivo, Estudo da reprodução e fecundidade do pargo, Lutjanus 
purpureus Poey, do norte e nordeste do Brasil. Arquivos de Ciências do Mar. 13, 109–
112 (1973). 
125. R. D. Evans, G. R. Russ, J. P. Kritzer, Batch fecundity of Lutjanus carponotatus 
(Lutjanidae) and implications of no-take marine reserves on the Great Barrier Reef, 
Australia. Coral Reefs 27, 179–189 (2008). doi:10.1007/s00338-007-0309-8 
126. T. C. V. Gesteira, C. A. S. Rocha, Estudo sobre a fecundidade do ariacó, Lutjanus synagris 
(Linnaeus), da costa do Estado do Ceará (Brasil). Arquivos de Ciências do Mar. 16, 19–
22 (1976). 
127. A. R. Mena-Loría, thesis, Centro de Investigación y de Estudios Avanzados del Instituto 
Politecnico Nacional, Instituto Politecnico Nacional, Mérida, Yucatá, Mexico (2009). 
128. C. B. Grimes, G. R. Huntsman, Reproductive biology of the vermilion snapper, 
Rhomboplites aurorubens, from North Carolina and South Carolina. Fish Bull. 78, 137–
146 (1980). 
129. J. L. Ross, J. V. Merriner, Reproductive biology of the blueline tilefish, Caulolatilus 
microps, off North Carolina and South Carolina. Fish Bull. 81, 553–568 (1983). 
130. J. F. Cerna, C. Oyarzún, Talla de primera madurez sexual y fecundidad parcial de la 
merluza común (Merluccius gayi, Guichenot 1848) del área de la pesquería industrial de 
la zona de Talcahuano, Chile. Investig. Mar. 26, 31–40 (1998). doi:10.4067/S0717-
71781998002600004 
43
131. G. J. Macchi, M. Pájaro, M. D. Ehrlich, “Fecundidad parcial y frecuencia reproductiva del 
efectivo patagónico de merluza (Merluccius hubbsi),” in INIDEP informe técnico (Mar 
del Plata: Instituto Nacional de Investigación y Desarrollo Pesquero, INIDEP, Mar del 
Plata, Argentina, 2006), pp. 1–13. 
132. S. Mehault, R. Domínguez-Petit, S. Cerviño, F. Saborido-Rey, Variability in total egg 
production and implications for management of the southern stock of European hake. 
Fish. Res. 104, 111–122 (2010). doi:10.1016/j.fishres.2010.03.019 
133. H. Kawase, Spawning behavior of the pygmy leatherjacket Brachaluteres jacksonianus 
(Monacanthidae) in southeastern Australia. Ichthyol. Res. 52, 194–197 (2005). 
doi:10.1007/s10228-005-0272-8 
134. H. Kawase, Reproductive ecology of the black reef leatherjacket, Eubalichthys bucephalus 
(Monacanthidae) in temperate Australia. Ichthyol. Res. 55, 294–298 (2008). 
doi:10.1007/s10228-007-0028-8 
135. G. W. Barlow, Spawning, eggs and larvae of the longnose filefish Oxymonacanthus 
longirostris, a monogamous coralivore. Environ. Biol. Fishes 20, 183–194 (1987). 
doi:10.1007/BF00004953 
136. H. Kawase, A. Nakazono, Embryonic and pre-larval development and otolith increments in 
two filefishes, Rudarius ercodes and Paramonacanthus japonicus (Monacanthidae). Jpn. 
J. Ichthyol. 41, 57–63 (1994). 
137. I. Mayer, S. E. Shackley, P. R. Witthames, Aspects of the reproductive biology of the bass, 
Dicentrarchus labrax L. II. Fecundity and pattern of oocyte development. J. Fish Biol. 
36, 141–148 (1990). doi:10.1111/j.1095-8649.1990.tb05590.x 
138. C. F. Hickling, A contribution to the natural history of the English grey mullets [Pisces, 
Mugilidae]. J. Mar. Biol. Assoc. U. K. 50, 609–633 (1970). 
doi:10.1017/S0025315400004914 
139. M. S. J. Greeley, D. R. Calder, R. A. Wallace, Oocyte growth and development in the 
striped mullet, Mugil cephalus, during seasonal ovarian recrudescence: Relationship to 
fecundity and size at maturity. Fish Bull. 85, 187–200 (1987). 
140. H. P. Das, The fecundity of grey mullet Mugil cephalus L. along the Goa Coast. Mahasagar 
10, 79–82 (1977). 
141. E. I. L. Silva, S. S. De Silva, Aspects of the biology of grey mullet, Mugil cephalus L., adult 
populations of a coastal lagoon in Sri Lanka. J. Fish Biol. 19, 1–10 (1981). 
doi:10.1111/j.1095-8649.1981.tb05806.x 
142. L. Alvarez-Lajonchere, The fecundity of mullet (Pisces, Mugilidae) from Cuban waters. J. 
Fish Biol. 21, 607–613 (1982). doi:10.1111/j.1095-8649.1982.tb02864.x 
143. E. H. Chan, T. E. Chua, Reproduction in the greenback grey mullet, Liza subviridis 
(Valenciennes, 1836). J. Fish Biol. 16, 505–519 (1980). doi:10.1111/j.1095-
8649.1980.tb03730.x 
144. N. B. Marshall, Egg size in Arctic, Antarctic and deep-sea fishes. Evolution 7, 328–341 
(1953). doi:10.1111/j.1558-5646.1953.tb00096.x 
44
145. E. S. Tereshchenko, The dynamics of population fecundity in Barents Sea capelin. ICES J. 
Mar. Sci. 59, 976–982 (2002). doi:10.1006/jmsc.2002.1257 
146. W. W. Morse, Reproduction of the summer flounder, Paralichthys dentatus (L.). J. Fish 
Biol. 19, 189–203 (1981). doi:10.1111/j.1095-8649.1981.tb05823.x 
147. G. J. Macchi, J. M. D. de Astarola, Ciclo reproductivo y fecundidad del lenguado, 
Paralichthys patagonicus Jordan, en Jordan y Goss 1889. Rev. Invest. Dessaro. Pesq. 10, 
73–83 (1996). 
148. S. M. Sogard, S. A. Berkeley, R. Fisher, Maternal effects in rockfishes Sebastes spp.: A 
comparison among species. Mar. Ecol. Prog. Ser. 360, 227–236 (2008). 
doi:10.3354/meps07468 
149. T. B. Bagenal, Annual variations in fish fecundity. J. Mar. Biol. Assoc. U. K. 36, 377–382 
(1957). doi:10.1017/S0025315400016866 
150. T. B. Bagenal, The breeding and fecundity of the Long Rough Dab Hippoglossoides 
platessoides (Fabr.) and the associated cycle in condition. J. Mar. Biol. Assoc. U. K. 36, 
339–375 (1957). doi:10.1017/S0025315400016854 
151. A. D. Rijnsdorp, Changes in fecundity of female North Sea plaice (Pleuronectes platessa 
L.) between three periods since 1900. ICES J. Mar. Sci. 48, 253–280 (1991). 
doi:10.1093/icesjms/48.3.253 
152. L. J. Buckley, A. S. Smigielski, T. A. Halavik, E. M. Caldarone, B. R. Burns, G. C. 
Laurence, Winter flounder Pseudopleuronectes americanus reproductive success. II. 
Effects of spawning time and female size on size, composition and viability of eggs and 
larvae. Mar. Ecol. Prog. Ser. 74, 125–135 (1991). doi:10.3354/meps074125 
153. A. C. Gundersen, O. S. Kjesbu, K. H. Nedreaas, A. Stene, Fecundity of Northeast Arctic 
Greenland halibut (Reinhardtius hippoglossoides). J. Northwest Atl. Fish. Sci. 25, 29–36 
(1999). doi:10.2960/J.v25.a3 
154. M. Arellano-Martínez, B. P. Ceballos-Vázquez, L. Hernández-Olalde, F. Galván-Magaña, 
Fecundity of Cortez angelfish Pomacanthus zonipectus (Teleostei: Pomacanthidae) off 
Espiritu Santo Island, Gulf of California, Mexico. Cienc. Mar. 32 (1A), 65–71 (2006). 
doi:10.7773/cm.v32i1.64 
155. W. C. Cummings, thesis, University of Miami (1968). 
156. K. D. Kavanagh, Larval brooding in the marine damselfish Acanthochromis polyacanthus 
(Pomacentridae) is correlated with highly divergent morphology, ontogeny and life-
history traits. Bull. Mar. Sci. 66, 321–337 (2000). 
157. T. Kokita, Potential latitudinal variation in egg size and number of a geographically 
widespread reef fish, revealed by common-environment experiments. Mar. Biol. 143, 
593–601 (2003). doi:10.1007/s00227-003-1104-x 
158. L. de L. G. Souza, S. Chellappa, H. de C. B. Gurgel, Biologia reprodutiva do peixe-donzela, 
Stegastes fuscus Cuvier, em arrecifes rochosos no nordeste do Brasil. Rev. Bras. Zool. 24, 
419–425 (2007). doi:10.1590/S0101-81752007000200020 
45
159. O. Marcus, K. Kusemiju, Some aspects of the reproductive biology of the clupeid Ilisha 
africana (Bloch) off the Lagos Coast, Nigeria. J. Fish Biol. 25, 679–689 (1984). 
doi:10.1111/j.1095-8649.1984.tb04914.x 
160. S. F. Hildebrand, L. E. Cable, “Development and life history of fourteen teleostean fishes at 
Beaufort, N. C.,” in Bulletin of the United States Bureaus of Fisheries (U.S. Government 
Printing Office, Washington, DC, 1930), pp. 383–488. 
161. G. R. Shepherd, C. B. Grimes, Reproduction of weakfish, Cynoscion regalis, in the New 
York Bight and evidence for geographically specific life history characteristics. Fish 
Bull. 82, 501–511 (1984). 
162. F. G. Azeredo, thesis, Universidade Federal do Paraná, Curitiba, Brazil, (2014). 
163. M. S. Love, G. E. McGowen, W. Westphal, R. J. Lavenberg, L. Martin, Aspects of the life 
history and fishery of the white croaker, Genyonemus uneatus (Sciaenidae), off 
California. Fish Bull. 82, 179–198 (1984). 
164. S. W. Ross, Reproduction of the banded drum, Larimus fasciatus, in North Carolina. Fish 
Bull. 82, 227–235 (1984). 
165. A. E. A. de M. Vazzoler, Sôbre a fecundidade e a desova da pescada-foguete. Boletim do 
Instituto Oceanográfico da Universidade de São Paulo 13, 117–122 (1963). 
166. A. E. A., de M. Vazzoler, Micropogon furnieri: Fecundidade e tipo de desova. Boletim do 
Instituto Oceanográfico da Universidade de São Paulo 18, 27–32 (1970). 
167. G. J. Macchi, M. E. Acha, C. A. Lasta, Desove y fecundidad de la corvina rubia 
Micropogonias furnieri Desmarest, 1823 del estuario del Río de la Plata, Argentina. Bol. 
Inst. Esp. Oceanogr. 12, 99–113 (1996). 
168. E. E. DeMartini, R. K. Fountain, Ovarian cycling frequency and batch fecundity in the 
queenfish, Seriphus politus: Attributes representative of serial spawning fishes. Fish Bull. 
79, 547–560 (1981). 
169. M. Inoue, Y. Iwasaki, M. Aoki, T. Kouhei, H. Nagaoka, Some notes on the artificial 
fertilization and rearing of larvae in frigate mackerel and yellowfin tuna, cultured with 
seawater-acclimatized Chlorella as a basic diet. J. Fac. Mar. Sci. Technol. Tokai Univ. 8, 
27–36 (1974). 
170. L. S. Emerson, M. G. Walker, P. R. Witthames, A stereological method for estimating fish 
fecundity. J. Fish Biol. 36, 721–730 (1990). doi:10.1111/j.1095-8649.1990.tb04326.x 
171. J. H. Finucane, L. A. Collins, H. A. Brusher, C. H. Saloman, Reproductive biology of king 
mackerel, Scomberomorus cavalla, from the southeastern United States. Fish Bull. 84, 
841–850 (1986). 
172. C. T. C. Ivo, Sobre a fecundidade da cavala, Scomberomorus cavalla (Cuvier), em águas 
costeiras do Estado do Ceará (Brasil). Arquivos de Ciências do Mar. 14, 87–89 (1974). 
173. T. C. V. Gesteira, Sobre a reprodução e fecundidade da serra, Scomberomorus maculatus 
(Mitchill), no Estado do Ceará. Arquivos de Ciências do Mar. 12, 117–122 (1972). 
46
174. J. H. Farley, A. J. Williams, S. D. Hoyle, C. R. Davies, S. J. Nicol, Reproductive dynamics 
and potential annual fecundity of South Pacific albacore tuna (Thunnus alalunga). PLOS 
ONE 8, e60577 (2013). doi:10.1371/journal.pone.0060577 Medline 
175. G. W. Bane, The Biology of the Yellowfin Tuna, (Bonnaterre) in the Gulf of Guinea (Cornell 
Univ. Press, 1963), vol. 2. 
176. R. E. J. Baglin, Reproductive biology of western Atlantic bluefin tuna. Fish Bull. 80, 121–
134 (1982). 
177. D. R. Gunderson, Population biology of Pacific Ocean perch, Sebastes alutus, stocks in the 
Washington-Queen Charlotte Sound region, and their response to fishing. Fish Bull. 75, 
369–403 (1977). 
178. L. A. Lisovenko, in Soviet Fisheries Investigations in the Northeast Pacific—Sovetskie 
rybokhozyaistvennye issledovaniya v severo-vostochnoi chasti Tikhogo okeana 
[Translated from Russian], P. A. Moiseev, Ed. (Israel Program for Scientific 
Translations, Jerusalem, Israel, 1965), pp. 162–169. 
179. A. C. DeLacy, C. R. Hitz, R. L. Dryfoos, “Maturation, gestation, and birth of rockfish 
(Sebasrodes) from Washington and adjacent waters,” in Fisheries Research Papers 
(Washington Department of Fisheries, WA, 1964), pp. 51–67. 
180. R. D. Stanley, A. R. Kronlund, Life history characteristics for silvergray rockfish (Sebastes 
brevispinis) in British Columbia waters and the implications for stock assessment and 
management. Fish Bull. 103, 670–684 (2005). 
181. P. M. Washington, R. Gowan, D. H. Ito, “A biological report on eight species of rockfish 
(Sebastes spp.) from Puget Sound, Washington” (Northwest and Alaska Fisheries Center 
Processed Report, National Oceanic and Atmospheric Administration, Seattle, WA, 
1978), pp. 1–50. 
182. M. S. Love, P. Morris, M. McCrae, R. Collins, “Life history aspects of 19 rockfish species 
(Scorpaenidae: Sebastes) from the Southern California Bight” (NOAA Technical Report 
NMFS, National Oceanic and Atmospheric Administration, Seattle, WA, 1978), pp. 1–
38. 
183. D. Benet, E. Dick, D. Pearson, “Life history aspects of greenspotted rockfish (Sebastes 
chlorostictus) from central California” (NOAA Technical Report NMFS, National 
Oceanic and Atmospheric Administration, Seattle, WA, USA, 2009), pp. 1–43. 
184. J. B. Phillips, “Life history studies on ten species of rockfish (genus Sebastodes),” in Fish 
Bulletin 126 (The Resources Agency of California, Department of Fish and Game, San 
Diego, CA, 1964), pp. 1–70. 
185. G. W. Boehlert, W. H. Barss, P. B. Lamberson, Fecundity of the widow rockfish, Sebastes 
entomelas, off the coast of Oregon. Fish Bull. 80, 881–884 (1982). 
186. V. A. Snytko, L. A. Borets, “Some data on fecundity of ocean perch in Vancouver-Oregon 
region” (Fisheries Research Board of Canada, Translation Series No. 2505, Department 
of the Environment, Fisheries Research Board of Canada Biological Station, Nanaimo, 
BC, Canada, 1973), pp. 249–251. 
47
187. S. J. Bobko, S. A. Berkeley, Maturity, ovarian cycle, fecundity, and age-specific parturition 
of black rockfish (Sebastes melanops). Fish Bull. 102, 418–429 (2004). 
188. S. G. Beyer, S. M. Sogard, C. J. Harvey, J. C. Field, Variability in rockfish (Sebastes spp.) 
fecundity: Species contrasts, maternal size effects, and spatial differences. Environ. Biol. 
Fishes 98, 81–100 (2015). doi:10.1007/s10641-014-0238-7 
189. J.-F. St-Pierre, Y. de Lafontaine, “Fecundity and reproduction characteristics of beaked 
redfish (Sebastes fasciatus and S. mentella) in the Gulf of St. Lawrence” (Canadian 
Technical Report of Fisheries and Aquatic Sciences, Division de la Productivitédes 
Océans Institut Maurice-Lamontagne Ministère des Pêches et Océans, Mont-Joli, 
Québec, Canada, 1995), pp. 1–32. 
190. F. Saborido-Rey, R. Domínguez-Petit, D. Garabana, Þ. Sigurðsson, Fecundity of Sebastes 
mentella and Sebastes norvegicus in the Irminger Sea and Icelandic waters. Cienc. Mar. 
41, 107–124 (2015). doi:10.7773/cm.v41i2.2500 
191. K. Schmidt, thesis, California State University, Monterey Bay, CA (2014). 
192. D. F. S. Raitt, W. B. Hall, On the fecundity of the redfish, Sebastes marinus (L.). ICES J. 
Mar. Sci. 31, 237–245 (1967). doi:10.1093/icesjms/31.2.237 
193. M. S. Love, K. Johnson, Aspects of the life histories of grass rockfish, Sebastes rastrelliger, 
and brown rockfish, S. auriculatus, from southern California. Fish Bull. 87, 100–109 
(1998). 
194. M. S. Love, W. V. Westphal, Growth, reproduction, and food habits of olive rockfish, 
Sebastes serranoldes, off central California. Fish Bull. 79, 533–545 (1981). 
195. K. V. Drevetnyak, A. I. Kluev, in 2005 ICES Annual Science Conference and Applications 
for Fisheries Science (International Council for the Exploration of the Seas, ICES, 
Aberdeen, UK, 2005), pp. 1–11. 
196. W. P. Nagelkerken, “Biology of the Graysby, Epinephelus cruentatus, of the coral reef of 
Curaçao,” in Studies on the Fauna of Curaçao and Other Caribbean Islands (The Hague, 
1979), pp. 1–118. 
197. A. Bouain, Y. Siau, Observations on the female reproductive cycle and fecundity of three 
species of groupers (Epinephelus) from the southeast Tunisian seashores. Mar. Biol. 73, 
211–220 (1983). doi:10.1007/BF00406890 
198. R. Abu-Hakima, Aspects of the reproductive biology of the grouper, Epinephelus tauvina 
(Forskål), in Kuwaiti waters. J. Fish Biol. 30, 213–222 (1987). doi:10.1111/j.1095-
8649.1987.tb05746.x 
199. V. Nóh-Quiñones, T. Brulé, C. Doralice, E. Peréz, in Proceedings of the 63rd Gulf and 
Caribbean Fisheries Institute (Gulf and Caribbean Fisheries Institute, Marathon, FL, 
2010), pp. 207–210. 
200. E. A. Wassef, H. A. A. Hady, Breeding biology of rabbitfish Siganus canaliculatus 
(Siganidae) in mid Arabian Gulf. Fish. Res. 33, 159–166 (1997). doi:10.1016/S0165-
7836(97)00075-1 
48
201. P. R. Witthames, M. Greer Walker, M. T. Dinis, C. L. Whiting, The geographical variation 
in the potential annual fecundity of dover sole Solea solea (L.) from European shelf 
waters during 1991. Neth. J. Sea Res. 34, 45–58 (1995). doi:10.1016/0077-
7579(95)90013-6 
202. R. Abu-Hakima, Some aspects of the reproductive biology of Acanthopagrus spp. (Family: 
Sparidae). J. Fish Biol. 25, 515–526 (1984). doi:10.1111/j.1095-8649.1984.tb04898.x 
203. P. Chavance, C. Flores-Coto, A. Sanchez-Iturbe, Early life history and adult biomass of sea 
bream in the Terminos Lagoon, southern Gulf of Mexico. Trans. Am. Fish. Soc. 113, 
166–177 (1984). doi:10.1577/1548-8659(1984)113<166:ELHAAB>2.0.CO;2 
204. S. Pati, On the maturation and spawning of Chinese pomfret Pampus chinensis (Euphrasen) 
from Orissa coast. Indian J. Fish. 26, 150–162 (1979). 
205. R. B. Silveira, thesis, Pontifícia Universidade Católica do Rio Grande do Sul, Porto Alegre, 
RS, Brazil (2005). 
206. J. D. Brown, thesis, University of Florida, FL (1972). 
207. W. Gladstone, M. Westoby, Growth and reproduction in Canthigaster valentini (Pisces, 
Tetraodontidae): A comparison of a toxic reef fish with other reef fishes. Environ. Biol. 
Fishes 21, 207–221 (1988). doi:10.1007/BF00004864 
208. J. A. Koslow, J. Bell, P. Virtue, D. C. Smith, Fecundity and its variability in orange roughy: 
Effects of population density, condition, egg size, and senescence. J. Fish Biol. 47, 1063–
1080 (1995). doi:10.1111/j.1095-8649.1995.tb06030.x 
209. A. R. S. Vieira, thesis, Universidade de Lisboa, Lisboa, Portugal (2007). 
210. N. Silverberg, H. M. Edenborn, G. Ouellet, P. Béland, Direct evidence of a mesopelagic 
fish, Melanostigma atlanticum (Zoarcidae) spawning within bottom sediments. Environ. 
Biol. Fishes 20, 195–202 (1987). doi:10.1007/BF00004954 
211. L. Smith, E. H. Atz, J. C. Tyler, Aspects of oral brooding in the cardinalfish Cheilodipterus 
affinis Poey (Apogonidae). Am. Mus. Novit. 2456, 1–11 (1971). 
212. D. A. Bengtson, R. C. Barkman, W. J. Berry, Relationships between maternal size, egg 
diameter, time of spawning season, temperature, and length at hatch of Atlantic 
silverside, Menidia menidia. J. Fish Biol. 31, 697–704 (1987). doi:10.1111/j.1095-
8649.1987.tb05272.x 
213. E. R. Waite, “Fishes,” in Australasian Antarctic Expedition 1911–1914. Scientific Reports. 
Series C—Zoology and Botany (Government Print, Adelaide, SA, Australia, 1916), pp. 
1–92. 
214. M. Akazaki, Biological studies on a dragonet, Synchiropus altivelis (Temminck et 
Schlegel). Jpn. J. Ichthyol. 5, 146–152 (1957). 
215. M. Demir, On the eggs and larvae of the Trachurus trachurus (L.) and Trachurus 
mediterraneus (Stdhnr.) from the Sea of Marmara and the Black Sea. Rapports et procès-
verbaux des réunions Comission Internationale pour LExploration Scientifique de la Mer 
Méditerranée. 16, 317–320 (1961). 
49
216. R. K. Pipe, P. Walker, The effect of temperature on development and hatching of scad, 
Trachurus trachurus L., eggs. J. Fish Biol. 31, 675–682 (1987). doi:10.1111/j.1095-
8649.1987.tb05270.x 
217. P. Pappenheim, Die fische der Deutschen Südpolar-Expedition 1901-1903. I. die fische der 
Antarktis und Subantarktis. Deutsche Südpolar Expedition 13, 161–182 (1912). 
218. R. J. Mansueti, Eggs, larvae, and young of the hickory shad, Alosa mediocris, with 
comments on its ecology in the estuary. Chesap. Sci. 3, 173–205 (1962). 
doi:10.2307/1350995 
219. W. F. Hettler, Description of eggs, larvae, and early juveniles of Gulf menhaden, Brevoortia 
patronus, and comparisons with Atlantic menhaden, B. tyrannus, and yellowfin 
menhaden, B. smithi. Fish Bull. 82, 85–95 (1984). 
220. J. D. de Ciechomski, Development of the larvae and variations in the size of the eggs of the 
argentine anchovy, Engraulis anchoita Hubbs and Marini. ICES J. Mar. Sci. 30, 281–290 
(1966). doi:10.1093/icesjms/30.3.281 
221. G. Marteinsdottir, K. W. Able, Geographic variation in egg size among populations of the 
mummichog, Fundulus heteroclitus (Pisces: Fundulidae). Copeia 1988, 471–478 (1988). 
doi:10.2307/1445889 
222. J. E. R. Cunningham, E. K. Balon, Early ontogeny of Adinia xenica (Pisces, 
Cyprinodontiformes): 1. The development of embryos in hiding. Environ. Biol. Fishes 
14, 115–166 (1985). doi:10.1007/BF00000821 
223. J. R. G. Hislop, The influence of maternal length and age on the size and weight of the eggs 
and the relative fecundity of the haddock, Melanogrammus aeglefinus, in British waters. 
J. Fish Biol. 32, 923–930 (1988). doi:10.1111/j.1095-8649.1988.tb05435.x 
224. C. Geevarghese, P. A. John, Maturation and spawning of a gobiid fish, Oligolepis 
acutipennis (Cuv. & Val.), from the south-west coast of India. J. Fish Biol. 23, 611–624 
(1983). doi:10.1111/j.1095-8649.1983.tb02941.x 
225. P. L. Colin, L. J. Bell, Aspects of the spawning of labrid and scarid fishes (Pisces: 
Labroidei) at Enewetak Atoll, Marshall Islands with notes on other families. Environ. 
Biol. Fishes 31, 229–260 (1991). doi:10.1007/BF00000690 
226. N. Raventos, S. Planes, Maternal size effects on early life traits of the temperate fish 
Symphodus roissali. Aquat. Biol. 4, 1–6 (2008). doi:10.3354/ab00086 
227. A. J. E. Lönnberg, “The fishes of the Swedish South Polar Expedition,” in 
Wissenschaftliche Ergebnisse der Schwedischen Südpolar-Expedition, 1901–1903 
(Lithographisches Institut des Generalstabs, Stockholm, 1905), vol. 5, pp. 1–69. 
228. C. M. Crawford, Development of eggs and larvae of the flounders Rhombosolea tapirina 
and Ammotretis rostratus (Pisces: Pleuronectidae). J. Fish Biol. 29, 325–334 (1986). 
doi:10.1111/j.1095-8649.1986.tb04949.x 
229. D. M. Buñag, Spawning habits of some Philippine tuna based on diameter measurements of 
the ovarian ova. Philippine J. Fisheries 4, 145–177 (1956). 
50
230. M. Demir, N. Demir, in Proceedings and Technical Papers (General Fisheries Council for 
the Mediterranean, 1961), pp. 213–218. 
231. S. Kumne, “A note on the fertilization of bigeye tuna, Parathunnus mebachi (Kishinouye),” 
Report of Nankai Regional Fisheries Research Laboratory (no. 15) (1962), pp. 79–84. 
232. L. R. Rivas, A preliminary report on the spawning of the western North Atlantic bluefin 
tuna (Thunnus thynnus) in the straits of Florida. Bull. Mar. Sci. Gulf Caribb. 4, 302–322 
(1954). 
233. W. G. Pearcy, Egg masses and early developmental stages of the scorpaenid fish, 
Sebastolobus. J. Fish. Res. Board Can. 19, 1169–1173 (1962). doi:10.1139/f62-079 
234. N. Devauchelle, J. Alexandre, N. Le Corre, Y. Letty, Spawning of sole (Solea solea) in 
capivity. Aquaculture 66, 125–147 (1987). doi:10.1016/0044-8486(87)90227-4 
235. I. Braga Goncalves, I. Ahnesjö, C. Kvarnemo, The relationship between female body size 
and egg size in pipefishes. J. Fish Biol. 78, 1847–1854 (2011). doi:10.1111/j.1095-
8649.2011.02984.x Medline 
236. J. R. Hunter, R. Leong, The spawning energetics of female northern anchovy, Engraulis 
mordax. Fish Bull. 79, 215–230 (1981). 
237. L. R. Castro, G. Claramunt, M. C. Krautz, A. Llanos-Rivera, P. Moreno, Egg trait variation 
in anchoveta Engraulis ringens: A maternal response to changing environmental 
conditions in contrasting spawning habitats. Mar. Ecol. Prog. Ser. 381, 237–248 (2009). 
doi:10.3354/meps07922 
238. S. S. Crawford, E. K. Balon, Alternative life histories of the genus Lucania: 3. An 
ecomorphological explanation of altricial (L. parva) and precocial (L. goodei) species. 
Environ. Biol. Fishes 41, 369–402 (1994). doi:10.1007/BF02197855 
239. J. R. G. Hislop, M. A. Bell, Observations on the size, dry weight and energy content of the 
eggs of some demersal fish species from British marine waters. J. Fish Biol. 31, 1–20 
(1987). doi:10.1111/j.1095-8649.1987.tb05209.x 
240. D. A. Fletcher, R. J. Wootton, A hierarchical response to differences in ration size in the 
reproductive performance of female three-spined sticklebacks. J. Fish Biol. 46, 657–668 
(1995). doi:10.1111/j.1095-8649.1995.tb01102.x 
241. N. Devauchelle, D. Coves, The characteristics of sea bass (Dicentrarchus labrax) eggs: 
Description, biochemical composition and hatching performances. Aquat. Living Resour. 
1, 223–230 (1988). doi:10.1051/alr:1988022 
242. R. N. Finn, H. J. Fyhn, M. S. Evjen, Respiration and nitrogen metabolism of Atlantic 
halibut eggs (Hippoglossus hippoglossus). Mar. Biol. 108, 11–19 (1991). 
doi:10.1007/BF01313466 
243. R. G. Bradford, R. L. Stephenson, Egg weight, fecundity, and gonad weight variability 
among Northwest Atlantic herring (Clupea harengus) populations. Can. J. Fish. Aquat. 
Sci. 49, 2045–2054 (1992). doi:10.1139/f92-227 
51
244. D. R. Robertson, S. E. Swearer, K. Kaufmann, E. B. Brothers, Settlement vs. environmental 
dynamics in a pelagic-spawning reef fish at Caribbean Panama. Ecol. Monogr. 69, 195–
218 (1999). doi:10.1890/0012-9615(1999)069[0195:SVEDIA]2.0.CO;2 
 
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Fish reproductive-energy output increases disproportionately with body size
Diego R. Barneche, D. Ross Robertson, Craig R. White and Dustin J. Marshall
DOI: 10.1126/science.aao6868
 (6389), 642-645.360Science 
, this issue p. 642Science
some fish stocks.
females. Management practices that ignore the value of large females could contribute to unexplained declines seen in 
 show that larger females are far more productive than the same weight's worth of smalleret al.incorrect. Barneche 
that suggest that one large female fish can be replaced by several smaller females. However, this assumption is
fecundity, regardless of the number of individuals involved. This assumption leads to fisheries management practices 
The theoretical relationship between reproduction and body size has assumed that total mass relates directly to
Big mamas matter for fish
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