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Is there any “magic” number behind the length at onset of sexual maturation
in marine fish and shellfish?

Article · September 2017

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Sergio Ragonese Patrizia Jereb

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Ragonese and Jereb (2017) - Is there any “magic” number behind the onset of sexual maturation? NTR-ITPP, sr77

Is there any “magic” number behind the length at onset

of sexual maturation in marine fish and shellfish?

Sergio Ragonese* and Patrizia Jereb**

*) Institute for Coastal Marine Environment (IAMC) of the National Research Council (CNR) of Mazara del Vallo, Via
Luigi Vaccara, 61, 91026 Mazara del Vallo (TP), Italy. E-mail corresponding Author: sergio.ragonese@iamc.cnr.it -
**) Italian National Institute for Environmental Protection and Research (ISPRA), Via Vitaliano Berancati 48/60,
00144 Rome, Italy.

Abstract – The dimensionless ratio(s) between length at maturity and maximum / asymptotic /
infinite length in fish and shellfish (also known as the 3rd Beverton and Holt’s Invariant) are
explored following the hypothesis that these values are distributed according some “magic
numbers”. Ten “magic” values were selected ranging between 0.11 (Liouville’s number) and 0.69
(Ln 2). The analysis of the invariants computed using data from fisheries literature suggest that the
suitable numbers (if any) should be searched in the ranges between 0.57 – 0.61 and 0.80 – 0.83.
However, there are evidences that current estimates are inflated by stock overexploitation that
determines reduced maximum lengths, only partially compensated by an analogous reduction in
length at maturity. A road map to re-estimate more properly the 3rd Beverton and Holt’s invariant is
proposed.

Key words: Life history invariant, Length at maturity, Maximum length, Asymptotic / Infinite length,
Magic numbers.

For bibliographic purpose this contribute should be quoted as:

Ragonese S. and P. Jereb (2017). Is there any “magic” number behind the onset of sexual maturation in
marine fish and shellfish? NTR-ITPP, sr77: 26 pp.

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Ragonese and Jereb (2017) - Is there any “magic” number behind the onset of sexual maturation? NTR-ITPP, sr77

Prologue
In 1963, four scientists won the Nobel prize for physics. Among these, the German physicist Maria
Goeppert-Mayer and the Hungarian physicist Edmund Wigner, both naturalized US citizens, who,
among other issues and working independently, developed the “shell” model for the atomic
nucleus. According to this model, there are given numbers of nucleons (protons and/or neutrons)
which make the nucleus particularly stable, so that even the increase/decrease in one single
nucleon makes the atomic structures less stable. These numbers, 2, 8, 20, 28, 50, 82 and 126, were
called “magic” numbers by Edmund Wigner.

Introduction

Notwithstanding the numerous proposals, conjectures, rules, theories and models produced
by fishery scientists (Berrigan and Charnov, 1994, Froese and Pauly, 2013, and other references
listed below) to describe the size at onset of sexual maturity (SOSM) in exploited marine fish and
shellfish, none of these was able to fully and satisfactory explain which mechanisms do drive this
complex phenomenon.

A few benchmarks, however, have been established.

The first is that size (both length and weight) and age at the onset of sexual maturity (OSM)
for a given stock reflect the plastic interaction among environmental conditions (especially
temperature and food availability) and the life history traits of that stock (especially the genetic
structure, growth, mortality, recruitment and fishing selectivity).

The second is that the OSM occurs at some discrete point of a trajectory, which may be
unique (Alm, 1959; Stearns and Crandall, 1984; Froese and Pauly, 2013; Figure 1) or develop
along multiple lines (i.e. given a minimum size and age several thresholds are achieved; cfr
Bernardo, 1983; Figure 2).

Not considering here those species showing determinate growth (Charnov et al., 1991;
Kozlowski et al., 2004), such as some crabs, in which the OSM coincides with a terminal moult, or
the “climax spawners” such salmons (Alverson and Carney, 1975), previous benchmarks mean that
in fish and shellfish showing an indeterminate growth (Kozlowski et al., 2004), although asymptotic
and a more (according to Bidder’s point of view; Bidder, 1932) or less (according to Comfort’s
point of view; Comfort, 1979) pseudo-infinite life span (Woodhead, 1998; Cailliet et al., 2001,
Ragonese, 2004; Vogt, 2012), the “relative size at maturity” (Charnov et al., 1991), i.e. the ratio
between the length at maturity (herein Lm for simplicity) and the maximum size achieved
(expressed as Lmax or L∞) can be expressed as a proportion ranging between 0.1-0.2 and 0.8-0.9
(figures too close to 0 or 1 being suspicious).

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Ragonese and Jereb (2017) - Is there any “magic” number behind the onset of sexual maturation? NTR-ITPP, sr77

Figure 1. – The benchmark of subsequent points along a single continuous line as explanation of
reproductive strategies in relation to the probability of survival (red curve) and the increase in body weight
(green curve). The blue curve indicates the expected reproductive output. Strategy A represents single-
spawners, such as salmons or eels. Strategy B represents multiple-spawners with parental investment, such as
live-bearers, nesters or guarders. Strategy C represents highly-fecund non-guarders such as cods, sardines or
tunas. The dotted horizontal lines indicate the necessary duration of the reproductive phase. From Froese and
Pauly, 2013.

Figure 2. - The benchmark of multiple possible lines around length and age thresholds along witch a stock
locates length at maturity. From Bernardo, 1993.

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However, yet early in the development of fisheries theory (Holt, 1958; Beverton and Holt,
1959), it was observed that the Lm value as a proportion of the asymptotic Length (L∞), in the
specific case consisting in the key parameter of the special von Bertalanffy Growth Function
(VBGF; Pauly, 1984), i.e. Lm/L∞, in fish consistently falls around a specific value, notably ≈ 0.66
(Beverton and Holt, 1959).

Thereafter, many other Authors have investigated this relationship and results obtained, both
empirically (Beddington and Kirkwood, 2005) and theoretically (Jensen, 1996), support what is
nowadays universally recognised as the 3rd Beverton & Holt’s Invariant (3BHI) (i.e. Lm/L∞ ≈ 0.66 –
0.67), also called Reproductive Load. However, data analysed already pointed out that the ratios
figures were distributed within a wider range of values (0.3–0.9; Holt, 1962).

The term “Invariant” (see also Life History Invariants; Charnov, 1993) is a general term
employed in fishery assessment studies to indicate dimensionless numbers (DLNs; Charnov et al.,
1991) which are considered of some utility in the assessment procedures, especially in case of
limited information about the investigated stock (Beddington and Kirkwood, 2005).

In other words, in spite of some severe criticisms (Nee et al., 2005), when studying stocks
which are very close taxonomically, it is assumed that the investigated invariant remain constant
(while it may slightly vary among different taxonomic groups).

Maturity related Invariants were also investigated for the relationships between age at sexual
maturity (Am) and the maximum age of a pristine stock (Ax), i.e., maximum age of the stock before
exploitation. In particular, Alverson and Carney (1975) proposed two values to describe these ratios
(0.25 and 0.38), but these figures were based on the assumption that Lm and the corresponding Am
were more or less coincident with the critical (optimum) length (Lcri) and age (Acri).

Lcri (also known as the Eumetric Length; Beddington and Kirkwood, 2005) in an unfished
stock is defined as the average size at which the specific growth rates and natural mortality rates are
equal (Ricker, 1945; Alverson and Carney, 1975).

This parameter, however, is more related to the so called length at “massive” (100%) sexual
maturation instead of to the more common length at 50% of maturity; as such, a Lcri /L∞ ratio value
of 0.75 is generally accepted in stock assessment studies and it was the first expression reported by
Holt (1958).

Further, although Lcri and Lm are highly correlated (r2 = 0.89–0.97), in small fishes Lcri may
be smaller-than or equal-to Lm, whereas in large fishes Lcri is usually larger than Lm (Froese and
Binohlan, 2000). In synthesis, the invariants computed using Lcri are expected to give different
results from those computed by using Lm.

Beside this problem, other maturity age/life span ratios can be found according to several
authors (Table 1).

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Ragonese and Jereb (2017) - Is there any “magic” number behind the onset of sexual maturation? NTR-ITPP, sr77

Table 1. Some examples of Age at sexual maturity (Am) / Life span (Ax) ratios.

Am/Ax ratios Taxon Reference

0.16 – 0.39 Bony fish Beverton, 1992

0.38 Gadiformes / Pleuronectiformes Beverton, 1992

0.57 Generic for marine animals Charnov, 1993

0.28 – 0.33 Fish Froese and Binohlan, 2000

0.38 Elasmobranchs Frisk et al., 2001

0.30 (a); 0.44 (b) a) Pelagic fish, b) Demersal fish Zhang and Megrey, 2006

0.23 - 0.31 Nothobranchius furzeri Terzibasi et al., 2007

0.11 Lutjanus campechanus Jarić and Gačić, 2012

0.06 Anoplopoma fimbria Jarić and Gačić, 2012

0.27 (a), 0.46 (b) a) Bony fish; b) Elasmobranchs Tsikliras and Stergiou, 2013

0.45 - 0.75 (an outlier at 0.20) Skates Barnett et al., 2013

Beside the wide range of values in Table 1 (0.06 - 0.75), it is to remark that:

a) most of the values fall between 0.4 and 0.6;

b) in Elasmobranchs (Frisk et al., 2001) and Gadiformes / Pleuronectiformes (Beverton,

1992) the same ratio occur (0.38), but

c) it is also noteworthy how this value is only slightly higher than the ratio’s range (0.23 -
0.31) obtained for the very short-living fish, Nothobranchius furzeri (life span =13 weeks; Terzibasi
et al., 2007).

In other words, fish seem to tend to achieve their sexual maturity at an age around 30–60%
of their maximum age no matter their absolute life span.

Although the existence of an Am/Ax invariant seems to be real, both age parameters present
high definition (Anthony, 1982; Cochran and Ellner, 1992; Hewitt and Hoenig, 2005) and
estimation (Barnett et al., 2013) problems. In particular, Barnett et al. (2013) highlighted that
theoretical life spans calculated from VBGF are likely overestimated and maximum observed ages
may underestimate the species’ true life span. Consequently, the age related invariants are no more
considered in this note.

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Ragonese and Jereb (2017) - Is there any “magic” number behind the onset of sexual maturation? NTR-ITPP, sr77

Coming back to the more familiar 3rd BHI (i,e. that based on L∞), it is worth recalling that
although Lm/L∞ values around 0.6–0.7 were confirmed by both theoretical and direct observations
(Charnov et al., 1991) even among terrestrial ectothermic animals such as lizards (0.73) and snakes
(0.64), evidences of a quite high variability of this invariants among the different fish and shellfish
stocks (from ≈ 0 up to ≈ 1; cfr Frisk et al., 2001) piled up.

This high variability, already evidenced in Beverton and Holt’s 1959 paper (cfr Figure 3),
reflects in part the poor precision of both Lm and L∞ parameters, due to many kinds of errors
(observational, sampling noise, methodological heterogeneity and biases). This imprecision is
additionally inflated by the combination of the two parameters to compute the ratio estimators; for
example, a simple 20% of error in Lm and L∞ estimates would determine an error up 40% in the
Lm/L∞ ratio.

Figure 3. - The original first Reproductive Load (i.e. Lm/ L∞) vs Longevity (life span) plot produced for fish
by Beverton and Holt (1959). Note the wide Lm/L∞ ratio range (0.3–0.9, with an outlier at 1.0) and the
asymmetric distribution with almost all long living species falling below the 0.6 value.

This said, fisheries scientists could consider two scenarios.

A) Sexual maturity related invariants are an illusion or computational artefacts, where the
observed coincidences with possible “magic numbers” (Mugnos, 2012) are casual. In this
scenario, the OSM occurs stock by stock always along a theoretical line (among multiple
possible lines), but with unclear predictable variability according to the general context. In
other words, length at maturity – maximum length relationships, although highly related to
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Ragonese and Jereb (2017) - Is there any “magic” number behind the onset of sexual maturation? NTR-ITPP, sr77

each other (Froese and Binohlan, 2002), should be estimated case by case without any
possible generalization.

B) Sexual maturity related invariants do exist, but the “magic value/s” is/are hidden given the
masking/camouflage effects of the heterogeneous and not comparable parameters used in
the computations. In synthesis, the conjecture behind this scenario can be expressed as “A
mathematical – geometric law might play a role in trigging the onset of maturity in marine
fish and shellfish”.

Assuming the B) scenario as the only one of interest in the present note, this implies that no
more than two “magic numbers” should be searched for.

In fact, considering 3 or more than 3 values, the corresponding distributions would result
statistically undistinguishable, especially for the proportions falling far from the midpoint 0.5,
characterised by high asymmetric / skewed distributions.

Aim of this note is to explore the possibility of figure out these “magic” numbers, which could
explain the relationship between the length at onset of sexual maturity and a maximum length
considered for the stock.

Material and Methods

A family of “magic” numbers was selected from the pertinent mathematical and geometrical
literature, according to a logical-guided process, i.e., trying to cover the expected range of the
hypothesized invariants values (Table 2).

These numbers were analysed on the base of the relationship between Lm and Lmax as reported in
the literature for a given stock; for simplicity, this ratio was referred to as the 3BHI. In particular,
Lmax may indicate the objective/empirical maximum ever recorded length as well as the subjective /
theoretical asymptotic length (also known as infinite; L∞). Where L∞ represents the parameter of the
simple / special VBGF (Pauly, 1984) if not otherwise specified.

Each length parameter and, consequently, the obtained relationships, present many definitional
and interpretative problems, as summarised in Table 3 and discussed below.

Whenever possible, the hidden normal components within the 3BHI frequency of distribution
were analysed by implementing the Bhattacharya and Normsep procedures as implemented in FiSat
(Gayanilo et al., 2005).

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Table 2. – The “magic” numbers derived from mathematic and geometry fields (in increasing order).

“Magic” number Remarks and references

0.11 Liouville’s number. Pickover, 2008

0.12 Champernoune’s number. Pickover, 2008

0.24 Copeland – Erdos’ number. Garcia del Cid, 2011

0.32 The inverse of pi Greco. Garcia del Cid, 2011

0.38 τ, Tau or φ, Phi. 1st Golden value. Garcia del Cid, 2011

0.57 Gamma / Euler – Mascheroni’s approximation. Garcia del Cid (2011)

0.62 2nd Golden value (Fibonacci’s series). Garcia del Cid, 2011

0.63 D = (Ln2/Ln3). Cantor’s number. Pickover, 2008

0.64 2/π. Ramanujan’s number. Garcia del Cid (2011)

0.69 Ln2. Pickover, 2008

Results and Discussion

By looking at the ten “magic numbers” selected (Table 2), it is worth remarking that no
magic number close to 1 was found in the literature.

Considering even a minimum of variability in both Lm and Lmax parameters listed in Table
3, it is possible to sort out 3 main intervals of magic numbers to use for close investigation: 1) early
maturity in relation to maximum length (0.11–0.30); 2) median maturity (0.30–0.60, and 3) late
maturity (above 0.60).

Two magic numbers, the so-called Golden ratios (GRs, also known as the “Golden mean”
or, even better, as the “Golden section”) deserve a special attention. They represent the only single
principle that offers two possible life styles to fish and shellfish. The GRs are two complementary
irrational (transcendent) numbers (Livio, 2002) properly defined for the first time by Euclid (367 -
283 a.C.). The Golden section is obtained by dividing a segment long a+b in two segments, a and b,
where the ratio between the entire segment and a is equal to the ratio between a and b, as described
in Figure 4.

The Golden section is known also for its peculiar mathematical characteristics; for example,
it is the only number the decimal section of which is equal to that of its squared number and to its
inverse (φ2= 2, 618033989 and 1/ φ = 0, 618033989).

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Table 3. - The basic metrics relevant for the invariants based on length at sexual maturity and maximum
length. Symbols according to Ragonese and Vitale, 2013.

Metric Denomination – Remarks / Main Reference

Lm.5 Length at (first) maturity or at 50% onset of sexual maturation. A too precocious stage to
distinguish the “mature” might result in an unrealistic low Lm (cfr Anon., 2016). No
consistent pattern about the variation of Lm with exploitation (Rochet, 2000a). A
reduction in high exploited stocks is commonly expected (Reznick, 1993; Hutchings,
2005; Marshall and Browman 2007; Sharpe and Hendry, 2009).

Lm1. Length at massive reproduction (≈100%). Associated to the critical age (Rikhter and
Efanov, 1976), but difficulty in choosing an objective value.

L∞ Theoretical length of an infinite aged fish or average length of the oldest fishes usually
referred to the special von Bertalanffy Growth Function (Ricker, 1979; Pauly, 1984). It
should be estimated when the stock is slight exploited. Difficult to estimate in exploited
stock. Extremely difficult to estimate (Bliss, 1970; Caddy, 1986) and normally affected
by subjective forcing resulting in lower values than expected. Usually considered
dependent on food availability (Ricker, 1979; Pauly, 1981; Francis 1990; 1995). It is
(improperly) considered equivalent to the maximum length (cfr Table 1 in Nee et al.,
2005).

Lxe - Lx Also Lmax. The single or mean of maximum lengths ever recorded (xe) or found (x) in
the exploited stock. Best defined as the length of the largest fish reported from a large
sample, possibly taken at a time when exploitation was still low. The Lxe is high reduced
just after the start of exploitation (Ricker, 1963). To not be confounded with Lλ (see
below).

∩Lcri Critical length (Ricker, 1945). Improperly named “optimal” or “optimum”. It should
strictly refer to the unexploited stock. Slight higher than Lm in large fish, slight lower
than Lm in small fish.

Lλ Length of ultimate significant contribute to the fishery at which adequate data are usable
for assessment (Beverton and Holt, 1957). Normally, it is much less than the maximum
or asymptotic length.

Lx vs Lm High log-linear correlation reported (Binohlan and Froese, 2009).

Lx vs L∞ Considered as equivalent, but to be reasonably, Lx must be computed on several

thousands of fish (Wetherall et al., 1987). L∞ ~ Lx/0.95 (Taylor 1958, at least in fish
smaller than 50cm, Pauly, 1984); L∞ > Lx (Ricker, 1975); no clear relationship (Mathews
and Samuel, 1990); high log linear correlated (Froese and Binohlan, 2000, but Lx/L∞
varies according the fish length intervals: 0.94 up 10cm, 0.97 up 100cm, 1.01 up 10m);
high simple linear correlation (Stergiou and Karachle, 2006).

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The original symbol used in the past to denote the Golden section (tau, τ, from tomé = cut)
was thereafter replaced by the symbol phi φ (or ) to honour Phidias (Athens 490– 430 a.C.).

Figure 4. – A simplified representation of the two (0.62 and 0.38) Golden ratios as “magic” numbers; φ (the
Golden section) = 1/0,618 = 0,618/0,382 = 1, 618033989.

The GRs find applications in several human activities (mathematics, geometry, architecture,
painting etc) and their occurrence was also found in the natural terrestrial and marine world (Livio,
2002; see for example the arrangement of leaves on a plant stem and the spirals / golden values
relationships in mollusc shells such as the Nautilus). An example of GR occurrence in fish is
presented in Figure 5.

Figure 5. – An example of magic number (as GR) in fish. The apex of pectoral fin (point B) individuates the
golden ratio point in the total length of the fish. From Snijders, 1993.

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It might be surprising to note the closeness of the 2nd GR (0.62) to the 3BHI (0.66) and the
coincidence of the 1st GR with the corresponding age based invariant (0.38) previously reported;
however, it is also to be remarked that other 3 magic numbers, D, 2/π and Ln(2), are closer to 0.66
than the 2nd GR.

Twenty-four 3BHIs central values (mean or median) were retrieved from the fisheries
literature (Appendix) resulting in an almost continuous sequence of values from 0.46 (Oreochromis
mossambicus; Binohlan and Froese, 2009) up 0.79 (Clupeiformes; Beverton, 1992). Only five ratios
occur twice (0.59, 0.62, 0.66, 0.70 and 0.77), supporting to some degree the consistency of 0.66 as
one of the magic number in OSM.

The graphic representations (Figures 6, 7 and 8) retrieved from fisheries literature (Tsikliras
and Stergiou, 2013; Froese and Pauly, 2013) may help some interpretation of this scattered
distribution. It is evident that, beside the problems which characterize the used parameters (cfr
Table 3), the 3BHIs are likely influenced by the systematic position of the investigated fish stocks
(bony vs selachians and Holocephali fish) and related reproductive behaviour (bearers or guarders
vs non-guarders).

As for the systematic issue (Figure 6), bony fish (Actinopterygii) show a lower median
invariant (0.59) than selachians (Elasmobranches; 0.70). The 0.77 value reported for Chimaera
(Holocephali) may be considered only indicative since it is based only on two species.

Figure 6. - Variability of the values of total length at maturity (Lm, cm) and maximum total length (Lmax, cm)
ratio per class (Actinopterygii, Elasmobranchii and Holocephali). The rectangular part of the plot extends
from the lower quartile to the upper quartile. The horizontal line within each box shows the location of the
mean. The whiskers extend from the box to the minimum and maximum values and circles indicate outliers.
From Tsikliras and Stergiou, 2013.

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Ragonese and Jereb (2017) - Is there any “magic” number behind the onset of sexual maturation? NTR-ITPP, sr77

As for the reproductive behaviour issue (Figures 7, 8), the median invariant looks significantly
higher in species showing some form of parental care than in non-guarders species, although with
different values according to the Authors.

Figure 7. - Total length at maturity (Lm, cm) to maximum total length (Lmax, cm) ratio for species providing
parental care (Yes) and those that do not (No). The rectangular part of the plot extends from the lower
quartile to the upper quartile. The horizontal line within each box shows the location of the mean. The
whiskers extend from the box to the minimum and maximum values and circles indicate outliers. From
Tsikliras and Stergiou, 2013.

Tsikliras and Stergiou (2013) reported 0.68 vs 0.59 for bearers or guarders compared to non-
guarders (Figure 7), whereas Froese and Pauly (2013), obtained 0.65 vs 0.50 for bearers or
guarders compared to non-guarders (Figure 8).

Beside the slight differences between the above resulting ranges, it is worth noting that the 0.65-
0.68 figures look very close to the expected 3BHI value (0.66) and are only slightly higher than the
2nd GR value. On the other side, the 0.50-0.59 values are lower than the 2nd GR, but quite higher
than the 1st GR (0.38).

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1.0

Length at first maturity 0.8

0.6

0.4

0.2

0.0
1-spawners bearer&guarders nonguarders

Figure 8. - Length at first maturity relative to asymptotic length for species that spawn only once in their life
time (1-spawners, 5 studies), bearers or guarders (49 studies), and non-guarders (178 studies). The horizontal
lines within the boxes present the median and the notched area the 95% confidence limits. The boxes contain
50% of the data and the extended lines indicate the spread of the data. The difference between bearers &
guarders and non guarders is significant. From Froese and Pauly, 2013.

Statistical differences were detected between the two groups (Figure 7), accepting which as
significant (i.e. high power of the test employed), only two hypothesis can be considered.

a) The previous reported invariants underestimate the true two magic numbers, which must be
searched for around 0.62–0.66 and 0.75 (the invariant based on the critical length),
respectively.
b) The previous reported invariants overestimate the true two magic numbers, which must be
searched for around 0.38 and 0.62 (i.e. the golden ratios).

To explore these hypotheses (i.e. only 2 magic numbers to be searched for, each of them with its
own distribution) a study of the frequency distribution of the invariants in the literature can be
carried out, in order to check the possibility of separating the expected two modal components.

Unfortunately, we were able to find only one invariant frequency distribution in the literature,
namely that in Kirkwood (1994), herein reported as Figure 9.

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Figure 9. - The corresponding reconstructed invariant frequencies distribution (starting from 0.2) are: 4.1,
6.9, 17.2, 21.3, 20.9, 16.6, 16.6, and 3.1. From Kirkwood, 1994.

Even an approximate look insight the distribution shown in Figure 9, suggests the existence of
at least two highly overlapping components, a feeling supported by the results of the attempted
deconvolution (Table 4; Figure 10).

Table 4. - An attempt to figure out the components of the Lm/L∞ ratio frequency distribution reported by
Kirkwood (1994) according to Bhattacharya's (A, B and C cases) and Normsep (D case) procedures, as
implemented in FiSat (Gayanilo et al., 2005). SD denotes the standard deviation of the mean. The component
discrimination to be objective, the Separation Index (SI) must be larger than 2. The corresponding plots are
presented in Figure 9.

Hypothesys Mean “magic” value SD SI

A - Only a single component A - 0.610 A - 0.165 ---

B – Two components, well separated B1 - 0.531 B1 - 0.120 ---

B2 - 0.770 B2 - 0.097 2.07

C – Two components, high overlapping C1 - 0.610 C1 - 0.165 ---

C2 - 0.800 C2 - 0.049 1.94

D – Automatic components definition D1 - 0.565 D1 - 0.149 ---

D2 - 0.831 D2 - 0.062 2.52

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Case Plot

Lm/L∞ 0.15 0.25 0.35 0.45 0.55 0.65 0.75 0.85 0.95

Figure 10. - Searching for modal components of the Lm /L∞ ratio frequency distribution reported by
Kirkwood (1994) according to Bhattacharya's (A, B and C cases) and Normsep (D case). See specifications
in Table 4 for details.

The unimodal fit (Case A) resulted in a median value almost coincident with the 2nd GR (0.61),
but with a standard deviation so large to make useless this ratio as an invariant.

Excluding the case C, where the separation index resulted lower than the critical limit of 2, two
components were detected by both a subjective (0.53 and 0.77; Case B) and objective/automatic
(0.56 and 0.83; case D) fit.

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According to these results, the a) hypothesis seems the most attractive, i.e. the magic numbers
might fall around 0.62 (the 2nd GR) and 0.75 (the invariant based on critical length).

However, there are some critical issues in the present analysis.

First, the “low” invariants in Tsikliras and Stergiou (2013) are based on the maximum length
(Lmax), whereas the “high” invariants in Kirkwood (1994) and Froese and Pauly (2013) were
computed using the asymptotic length (L∞).

In addition, it remains unclear why even using the same parameters (as in Kirkwood, 1994) the
computed ratios underestimate the first magic number (0.56 vs 0.62) and overestimate the second
magic number (0.83 vs 0.75).

In other words, there are some problems in the used parameters.

Several doubts refer to the suitability of the asymptotic/infinite length (L∞) for the invariant
computations (cfr Table 3). The first problem concerns the source of the parameter, i.e., the growth
model to which it is referred. Mostly invariants are implicitly based on L∞ estimated within the so-
called simple – special (Pauly, 1984) 3 parameters VBGF (Brody - von Bertalanffy growth
function; Ricker, 1979; Pauly, 1984). The suitability of the special VBGF has been strongly
criticised by many Authors (for example, Kozlowski et al., 2004; Charnov, 2008) and there are
evidences that published L∞ are quite imprecise and high subjective estimations.

The basic problems hampering a proper estimation of L∞ are the quick elimination of large sized
specimens just after the beginning of the exploitation (Ricker, 1963) and the rarefaction of adult
specimens; consequently, L∞ is variable reduced according to the geographical fishing patterns,
resulting in a very high dispersion of L∞ even in the same species (cfr Hake in Ragonese et al.,
2012).

However, also the maximum length is a problematic parameter. The maximum ever recorded
length is rarely available in historically overfished stocks (such as the Mediterranean cases) since
the lack of documented measures at the beginning of the fisheries, whereas the current maximum
length would require a very large sample (i.e. thousands of specimens) to compensate for the rarity
and high dispersion of the few occurring largest specimens. Further, there is no general agreement
about its definition (Table 3). There are evidences that a mean length estimated on the base of a
variable number of largest specimens (cfr Probst et al., 2013) should be preferable to the classic
single maximum length (Hoenig’s suggestion). Unfortunately, usually only the single maximum
value is available in published papers.

In conclusion, in highly exploited stocks, what is estimable is the maximum length at significant
statistical contribution to the fishery, i.e. the Lλ proposed by Beverton and Holt (1957; cfr Table 3),
that is by definition quite lower than Lmax and often used by scientists to calibrate the L∞. The third
parameter involved in the 3BHI is the length at onset of sexual maturity.

Conventionally, it is defined as the length at which 50% of the stock can be considered
“mature” (herein Lm.5) according to the application of a predefined macroscopic multi-stages scale
of gonads status classification (cfr the scales adopted within the MEDITS program; Anon. 2016)
modelled by a logistic (antisymmetric) curve.
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Ragonese and Jereb (2017) - Is there any “magic” number behind the onset of sexual maturation? NTR-ITPP, sr77

Beside the rare case of Lm.5 reduction in some fish such Lamprey (cfr Froese and Binohlan,
2000), the most critical sides of this parameter are:

1) it should be estimated from samples gathered during the peak of sexual activity of the stock
(and in most papers this requirement is not specified); 2) the stage used to distinguish
immature/mature specimens should reflect the mid-point of the gonads development (and in most
cases a very precocious stage is selected; cfr Anon., 2016); 3) the length at about 100% of maturity
should be considered the most proper parameter for the invariant since is more related to the critical
length (cfr Rikhter and Efanov, 1976; Ragonese and Bianchini, 2014); and 4) whichever the choice
between 50 or 100% maturity, there is great uncertainty about the invariance of this parameter in
function of an increased fishing mortality (fisheries induced evolution; Conover, 2002; Jørgensen et
al., 2007).

All the previous problems combined would determine an underestimation of the length at
maturity that should partially compensate for the reduced maximum / asymptotic length previously
exposed. The Lm at ≈ 100% could be arbitrarily approximated (Froese and Binohlan, 2000)
considering the reproductive load (Beverton, 1992) as Lm1. = Lm.5 +0·25*(L∞ – Lm.5), but this
procedure will introduce a circularity problem if L∞ will be used in the successive computations.

The final source of confusion to be recalled consists in the sex related differences in both
maximum length achieved and size at onset of sexual maturity; sexes should be treated separately
within the invariant estimation (cfr Tsikliras and Stergiou, 2013).

Conclusions

There is no general agreement about the suitability of the use/applications of invariants in

fishery science. Length-based invariants, in particular, are not considered a valuable field of
research by many relevant and authoritative Authors. Daniel Pauly, for example (personal
communications; but see also Pauly, 2010), believes that the reproductive load expressed as a
fraction of length (i.e. Lm/L∞) is not close to the underlying trigger for spawning, but that the
fraction of body weight at spawning relative to maximum body weight is. Indeed, this author
considers length as a measure of size not metabolically relevant.

Severe criticisms are also related to the “magic” numbers; Pommersheim et al., 2010, for
example, have argued that many apparent manifestations of the Golden ratio in nature, especially
concerning animal dimensions, are fictitious.

Notwithstanding the difficulties in dealing with highly imprecise ratios estimators and the
general scepticism about the existence of length related invariants and corresponding “magic”
values, published data suggest that at least close taxonomic stocks may evolve some reproductive
strategy implying the convergence of Lm/Lmax ratio towards a well determined “magic” value.

Golden ratios, in particular, seem the more suggestive candidates since they suit the two
basic fish reproductive strategies: achieve maturity relatively “early” or relatively “late” within a
given life span. According to this dichotomy, while Clupeiformes, for example, adopted the option
“First grow, then reproduce” (i.e. Lm/L∞  0.7), Gadiformes and Perciformes go the other way
17
Ragonese and Jereb (2017) - Is there any “magic” number behind the onset of sexual maturation? NTR-ITPP, sr77

round “Early maturity at small size relative to adult size” (i.e. Lm/L∞  0.4) and an intermediate
strategy is shown by Pleuronectiformes (see Rochet, 2000b).

It is worth remarking that the option chosen seem independent from the life span extension,
although the longer the life span is, the higher the convergence towards the 1 st Golden ratio (0.38)
would be expected, as suggested by the plot in Beverton and Holt 1959’s paper (Figure 3).

However, the above assumption is not so strictly applicable to the different fish populations;
for example (see Table 3 for the symbols), early maturity (i.e. Lm1./L∞ of 0.51 with a range of 0.25
– 0.77, Ragonese et al., 2012) can be found in the Mediterranean Hake (life span from 20-25 years;
Vitale et al., 2016), whereas late maturity occur in very long-living species (such as Sebastes;
Beverton, 1992), in medium-long living ones (such as sharks, 0.73, Frisk et al., 2001) and even in
annual (1 y of life span) semelparous species such as Aphia minuta (Lm/Lmax = 0.63).

Consequently, the suggestion to implement studies and research focused to improve our
knowledge on length-at-maturity based invariant seems worth of attention. To this goal, great
attention in selecting the most adequate parameters to be investigated is a basic requirement, to help
reducing the confounding effects of methodological heterogeneity and imprecision of the estimates.

A plausible methodological road map might be.

a) Sort out the Large Marine Ecosystem (LME) of reference, to begin with the Mediterranean
Sea, as a case study.
b) For each fish and shellfish stock, collect data (preferably by sex) on the documented single
Lxe recorded, both before the onset of modern exploitation and under the current exploitation
pattern.
c) For each fish and shellfish stock, collect the length value at the onset of sexual maturity,
preferably that corresponding to the 75 – 100% of maturity (i.e. the so called length at
“massive” maturation) both before the modern exploitation and under the current
exploitation pattern.
d) Pay much attention in avoiding the risk of circularity (i.e. using estimates derived by other
estimates).
e) Represent the derived ratios by box plots, both overall and by taxonomic groups (bony fish,
selachians, crustaceans, shrimps etc) in order to investigate the systematic differentials (if
any) and the degree of symmetry.
f) In case of high asymmetric ratios, use data transformation (such as arcsine).
g) Build up the frequency of distribution of the derived ratios and attempt to discriminate the
modal components by performing a mixture-solving software such as Normsep or Mix.
h) Compare the correspondence between the modal ratio of each component with the “magic”
numbers in order to check the existence of a significant statistical affinity.

Such analysis should be carried out for as many as possible LMEs in order to finally give an
answer to the question raised at the beginning of the present note:

Is there any “magic” number behind the length at onset of sexual maturation in fish and
shellfish?
18
Ragonese and Jereb (2017) - Is there any “magic” number behind the onset of sexual maturation? NTR-ITPP, sr77

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APPENDIX – Length at maturity (Lm) - critical length (Lcri) vs maximum / infinite / asymptotic length (Lmax;
L∞) reported in fisheries literature. See Table 2 for parameters specifications. Min: Minimum; Max:
Maximum; C value: Central value (mean or median); sd: standard deviation.

Taxon Min Max C value Remarks and references

Fish 0.30 0.90 0.66 Real estimates. Lm, L∞. Beverton and Holt (1959)

Females sharks 0.64 0.92 --- 16 species. Holden (1972) in Tsikliras and Stergiou, 2013 (as
Lm/Lmax).

Fish --- --- 0.75 63 species. Theoretical derivation; A0 = 0; M = K. Lcri, L∞.

Alverson and Carney (1975). The same 0.75 figure, but as
Lm/L∞, was also reported by Charnov et al. (1991).

Carcharhinus 0.50 0.85 --- Garrick (1982) in Tsikliras and Stergiou, 2013 (as Lm/Lmax).
males

Carcharhinus 0.60 0.90 --- Garrick 1982 in Tsikliras and Stergiou, 2013 (as Lm/Lmax).
females

Crustaceans --- --- 0.56 Five species of Pandalids. Lm, L∞. Charnov et al. (1991).

Fish (Antarctic 0.55 0.80 --- Kock and Kellermann (1991) in Tsikliras and Stergiou, 2013 (as
notothenioid) Lm/Lmax).

Flatfish, males 0.40 0.61 0.47 Real estimates. Lm, L∞. Beverton (1992).

Flatfish, females 0.45 0.62 0.52 Real estimates. Lm, L∞. Beverton (1992).

Gadiformes 0.45 0.78 0.61 Real estimates. Lm, L∞. Beverton (1992).

Sebastes - 0.64 0.78 0.72 Real estimates. Lm, L∞. Beverton (1992).
Atlantic

Sebastes - 0.66 0.82 0.78 Real estimates. Lm, L∞. Beverton (1992).
Atlantic

Clupeiformes 0.70 0.88 0.79 Real estimates. Lm, L∞. Beverton (1992).

Fish --- --- 0.66 Assuming that fish of a stock mature when their fecundity
function f(a) at age a takes its maximum value. Lm, L∞. Jensen
(1996).

Modelled 0.40 0.77 ---- With indeterminate growth. Kozlowski (1996) in Tsikliras and
animals Stergiou, 2013 (as Lm/Lmax).

Fish --- --- 0.714 Lm, Lmax. Pauly (1998).

Fish --- --- 0.62 Lcri, L∞. Froese and Binohlan (2000).

Selachians 0.19 1.00 0.73 sd = ± 0.022. Real estimates. Lcri, L∞. Frisk et al. (2001).

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Ragonese and Jereb (2017) - Is there any “magic” number behind the onset of sexual maturation? NTR-ITPP, sr77

Skates and Rays 0.67 0.85 0.77 Lm, Lmax. Reynolds et al. (2001). Based on Walker and
Hislop’s data.

Fish --- --- 0.62 141 stocks. Lcri, L∞. CV = 24 %. Cubillos (2003).

Fish 0.38 0.58 0.46 Oreochromis mossambicus. Lm, L∞. Geometric mean. Binohlan
and Froese (2009).

Fish 0.5 0.9 0.7 Species that spawn only once in their lifetime. 5 studies. Lm,
L∞. Median value. Froese and Pauly (2013).

Fish 0.4 0.9 0.65 Bearers or guarders. 49 studies. Lm, L∞. Median value. Froese
and Pauly (2013).

Fish 0.25 0.9 0.50 Non guarders. 178 studies. Lm, L∞. Median value. Froese and
Pauly (2013).

Fish 0.2 0.9 0.59 sd ± 0.122. 471 cases. Lm /Lmax. Tsikliras and Stergiou (2013).
(Actinopterygii)

Fish 0.4 0.9 0.70 sd ± 0.132. 92 cases. Lm /Lmax. Tsikliras and Stergiou (2013).
(Elasmobranchii)

Fish --- --- 0.77 sd ± 0.077. 2 cases. Lm /Lmax. Tsikliras and Stergiou (2013).
(Holocephali)

Fish (guarders, 0.25 0.90 0.68 sd ± 0.141. 111 cases. Lm /Lmax. Tsikliras and Stergiou (2013).
bearers, nesters)

Fish (non- 0.4 0.9 0.59 sd ± 0.123. 454 cases. Lm /Lmax. Tsikliras and Stergiou (2013).
guarders)

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 Ragonese and Jereb (2017) - Is there any “magic” number behind the onset of sexual maturation? NTR-ITPP, sr77

PRESENTAZIONE
Non mancano, in Italia e nel mondo, riviste scientifiche specializzate per le tematiche di cui si interessa e si
interesserà la sede secondaria di Mazara del Vallo dell’Istituto per lo studio dell’Ambiente Marino e Costiero
(IAMC) del Consiglio Nazionale delle Ricerche (CNR) e ad esse hanno chiesto e chiederanno ospitalità i
ricercatori della sede. Tuttavia, nel corso della sua attività, la sede IAMC CNR di Mazara produce anche, sia
direttamente che attraverso collaborazioni esterne, note tecniche e prime versioni di lavori scientifici non
ancora pronti per la stampa, ma che talvolta sembra opportuno e urgente diffondere negli ambienti scientifici
interessati. Diversi anni fa, prima della riforma del CNR quando la sede di Mazara era l’Istituto per la
Tecnologia della Pesca e del Pescato (ITPP – CNR), il Direttore Dino Levi istituì la pubblicazione
denominata Note Tecniche e Reprints (NTR – ITPP). Pubblicare nella serie degli NTR-ITPP, non era
necessariamente alternativo alla pubblicazione su riviste periodiche, internazionalmente riconosciute e con
autorevoli comitati editoriali, ma consentiva di diffondere in tempi brevi risultati interessanti ed importanti
che altrimenti avrebbero atteso tempi molto più lunghi per essere divulgati. NTR-ITPP inoltre, come dice il
nome, si proponeva come mezzo di rapida diffusione di reprints di lavori già pubblicati su Riviste
Scientifiche, a seguito di opportuna autorizzazione. NTR-ITPP infine, pur collocandosi, tipicamente, nella
cosiddetta "letteratura grigia", è a tutti gli effetti pubblicazione scientifica ufficialmente registrata e tutelata
dalla legge nazionale ed internazionale. In attesa che lo IAMC CNR si fornisca di un’analoga pubblicazione,
la sede secondaria di Mazara utilizzerà questo mezzo, mantenendo, per rispetto del fondatore Dino Levi e per
la memoria storica della prima istituzione del CNR a Mazara, il vecchio acronimo, pur specificando che si
tratta di una pubblicazione della sede secondaria IAMC CNR di Mazara del Vallo.

NTR- ITPP, Note Tecniche e Reprints della sede secondaria IAMC CNR di Mazara.
Reg. Tribunale di Marsala n° 76/1/88 del 9 Gennaio 1988 ISSN 1120-1894
Indirizzo: Via Luigi Vaccara, 61, 91026, Mazara del Vallo (TP), Italia
Telefoni: 0923/948966; 934116; Fax: 0923/906634.

PRESENTATION
In Italy and around the world there are many specialist magazines which publish papers and documents on
the same subjects as those on which the territorial branch of the Institute for Coastal Marine Environment
(ICME) of the National Research Council (NRC) in Mazara del Vallo carried out, and will continue to carry
out research. The researchers of the Branch of Mazara have always availed themselves of the hospitality
offered by these publications, and this collaboration will not cease. However, the Branch of Mazara often
produce technical notes and preliminary versions of studies not ready yet for definitive publication, but of
such a nature and importance to make it adequate and necessary their quick publication and diffusion among
the interested scientific community. Several years ago, well before the reorganisation of the NRC, when the
Institute of Technology of Fisheries and Fisheries products (ITPP) of Mazara was joined with other Institutes
to create the ICME, the Director of the Institute of Mazara del Vallo, Dino Levi, founded a publication
named NTR-ITPP, i.e. Technical Notes and Reprints of the ITPP. The NTR-ITPP series, should not be
looked at as an alternative to, or as a competitor of other periodicals which, though internationally
recognised and disposing of prestigious editing committees, unfortunately work/operate too slowly to be
useful in given circumstances. The NTR- ITPP series also present/disseminate reprints of already published
literature, given the necessary authorisations. Though it may be considered a typical example of the so-called
"grey literature" (it participates in the SIGLE program), NTR-ITPP is, to all effects, an officially registered
scientific publication, subject to the relative national and international legislation.

NTR- ITPP, Note Tecniche e Reprints of the ICME-NRC Branch of Mazara

Registration: Tribunale di Marsala n° 76/1/88 - 9th January 1988 - ISSN 1120-1894
Address: Via Luigi Vaccara, 61, 91026, Mazara del Vallo (TP), Italy
Phone: +39 0923/948966; +39 934116; Fax: +39 0923/906634.

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