Regional Studies in Marine Science 39 (2020) 101468

Contents lists available at ScienceDirect

Regional Studies in Marine Science

journal homepage: www.elsevier.com/locate/rsma

Morphometric relationships and non-parametric,

length–frequency-based growth parameters and natural mortality for
common marine forage fish in Florida waters (USA)
∗
Joseph Munyandorero , Dustin Addis
Fish and Wildlife Research Institute, Florida Fish and Wildlife Conservation Commission, 100 8th Avenue SE, St. Petersburg, FL 33701, USA

article info a b s t r a c t

Article history: Life-history parameters are necessary in population and ecosystem analyses. But they are not always
Received 2 June 2020 available, especially for species that, justifiably or not, are of low priority, such as the forage fish
Received in revised form 11 September 2020 of Florida, USA. When these parameters are unavailable for analyses, they are often borrowed from
Accepted 14 September 2020
species of other areas. To keep the borrowing of life-history parameters to a minimum for Florida
Available online 15 September 2020
forage fish, this study is devoted to developing the morphometric relationships, growth parameters,
Keywords: and natural mortality rates for forage fish species for which size data had been collected in inshore
Length frequencies surveys that have been conducted during 1996–2017 by the Fishery Independent Monitoring program
Conversion coefficients of the Florida Fish and Wildlife Conservation Commission’s Fish and Wildlife Research Institute. The
Growth parameters parameters developed can be used in single-species stock assessments and in marine ecosystem
Natural mortality modeling.
Forage fish © 2020 Elsevier B.V. All rights reserved.
Florida

1. Introduction species-specific stock assessments, whether the species is directly

Florida (USA) waters of the Atlantic Ocean, including the Gulf
of Mexico, have a diverse ichthyofauna. This ichthyofauna can be
subdivided into two broad categories based on life-history data
and parameters required in planning natural resources conserva-
tion. The first category comprises a few fish species for which
size, age, and reproduction data have been routinely collected,
and the estimated parameters based on those data have been reg-
ularly updated for federal and Florida-specific stock assessments
(e.g., Crabtree and Bullock, 1998; Bortone, 2003; White and
Palmer, 2004; Brown-Peterson et al., 2009; Adams et al., 2018;
Herdter et al., 2019; Young et al., in press). The second category
comprises an assortment of primarily forage fish (small, short-
lived, and low-trophic-level species; Ohs et al., 2013) for which
only length and weight data at best are available. Overall, life-
history data on forage fish in Florida are lacking, which hinders
the assessment of their status. Yet those fish can form significant
bycatch or discards, have often been directly targeted as bait used
to attract more valuable, predatory fish (hence their other name,
bait fish; Ohs et al., 2013), and may be the primary prey of marine
predators (Hilborn et al., 2017; Koehn et al., 2017).
Because forage fish are so economically and ecologically im-
portant, their status should be determined separately through
∗ Corresponding author.
E-mail address: Joseph.Munyandorero@MyFwc.com (J. Munyandorero).
targeted or is bycatch, or jointly in analyses aiming at ecosystem
management (e.g., Walters et al., 2008; Grüss et al., 2016).
Such status determinations require basic life-history parameters
on growth, morphometric relationships (i.e., equations whose
coefficients can be used to convert size types to one another),
and natural mortality. When such parameters are not available,
researchers often borrow them from the literature on similar
species from other areas and substitute them in analyses.
The present study addresses the estimation of morphometric
relationships (length–length and weight–length), growth param-
eters, and natural mortality of Florida’s forage fish, because those
life-history variables may be required in species-specific stock as-
sessments or ecosystem modeling of Florida waters. All estimates
are based on length and weight measurements for fish that were
sampled by the Fishery Independent Monitoring program (FIM)
of the Florida Fish and Wildlife Conservation Commission’s (FWC)
Fish and Wildlife Research Institute (FWRI).
2. Materials and methods
2.1. Survey method, areas, gear types, and data
During 1996–2017, the FIM program used monthly stratified
random surveys to collect continuously (typically each day) data
on animals inhabiting bays and estuaries of Florida (Fig. 1). The
types of sampling gear commonly used were a 21.3-m center-bag
seine, a 6.1-m otter trawl, and a 183-m haul seine, designed to

https://doi.org/10.1016/j.rsma.2020.101468
2352-4855/© 2020 Elsevier B.V. All rights reserved.
J. Munyandorero and D. Addis
Table 1
Marine forage fish of Florida (USA) studied (the scientific names and authorities
came from www.fishbase.org).
Common name Scientific name
Atlantic croaker Micropogonias undulates (Linnaeus, 1766)
Atlantic thread herring Opisthonema oglinum (Lesueur, 1818)
Bay anchovy Anchoa mitchilli (Valenciennes, 1848)
Pigfish Orthopristis chrysoptera (Linnaeus, 1766)
Pinfish Lagodon rhomboides (Linnaeus, 1766)
Sand perch Diplectrum formosum (Linnaeus, 1766)
Scaled sardinea Harengula jaguana (Poey, 1865)
Silver mulletb Mugil curema (Valenciennes, 1836)
Silver perch Bairdiella chrysura (Lacepède, 1802)
Spanish sardine Sardinella aurita (Valenciennes, 1847)
Spot croakerc Leiostomus xanthurus (Lacepède, 1802)
Striped mojarra Eugerres plumieri (Cuvier, 1830)
Striped mullet Mugil cephalus (Linnaeus, 1758)
a the usual VBGF Lt = L∞ 1 − e−K (t −t0 ) where Lt is FL at age
Known elsewhere as round sardinella and scaled herring.
b
Also called white mullet in Florida.
c
Simply called spot on the U.S. Atlantic coast and Gulf of Mexico.
collect, respectively, juvenile and subadult fishes in shallow areas
(<1.8 m); juvenile, subadult, and adult fishes in deep waters (1–
7.6 m); and subadult and adult fishes in shallow waters (< 2.5
m). In these sample catches, this study is focused on forage fish
(Table 1). Data recorded for forage fish consisted of length (mm)
and total wet weight (g).
During a sampling trip, a maximum of the first 20 individuals
of a forage fish collected by set of a gear type were measured
for standard length (SL, in mm; SL is the length of a fish from
the most anterior part of the body to the end of the hypural
plate) and distributed across length bins. When more than 20
fish were caught, length frequencies of the 20 fish were raised
to the total number caught to estimate the sample catch length
frequency. In the present study, length and weight measurements
were combined by coast and month across gear types and across
years. Standard lengths were converted into fork lengths (FL, the
length of a fish from the most anterior part of the body to the tip
of the medial caudal fin ray) because fishery analyses commonly
use parameters based on FL.
2.2. Morphometric relationships
The morphometric relationships were equations for convert-
ing SL to FL, SL to total length (TL, the length of a fish from the
most anterior part of the body to the tip of the longest caudal
fin ray when the lobes are compressed dorsoventrally), and TL to
FL, as well as SL, FL, and TL to wet weight. Lengths were in cm
and weights in g. Regression analyses were used to estimate the
parameters of various morphometric relationships (Tables 2–7).
2.3. Growth parameters, curves, and natural mortality
Bay anchovies were assigned to 0.5-cm length bins, Atlantic
croaker and silver mullet to 2-cm length bins, and the remain-
ing species to 1-cm length bins. These length bins were es-
sentially chosen considering that a species was more or less
small- or medium-sized; they were checked post-hoc against, and
found to be overall consistent with, the bin sizes that would be
calculated using the empirical equation of Wang et al. (2020),
.6
i.e., length bin = 0.3L0max , where Lmax is the maximum observed
length.
The monthly length frequencies obtained for each species
formed the basis for estimating the parameters of the von Berta-
lanffy growth function (VBGF) using the R package TropFishR,
version 1.6.2 (Mildenberger et al., 2017). TropFishR was designed
Regional Studies in Marine Science 39 (2020) 101468
to implement different routines of the software called FiSAT II
(Gayanilo et al., 1996), which uses length–frequency (LFQ) data
to assess data-poor stocks. In addition to the traditional version
Code
of the electronic length–frequency analysis (ELEFAN) used to
ATC
ATH
estimate the VBGF parameters, TropFishR includes optimization
BAN techniques for VBGF fitting (Taylor and Mildenberger, 2017).
PGF The derivation of the VBGF parameters from LFQ data was
PNF based on the TropFishR function ELEFAN_GA. Like the traditional
SNP
SCS ELEFAN method, ELEFAN_GA is a nonparametric procedure, but
SLM ELEFAN_GA uses a genetic (i.e., stochastic search) algorithm for
SLP solving optimization problems. Because of the stochastic nature
SPS of the function ELEFAN_GA, each run of TropFishR on the same
SPT
SPM LFQ data results in a unique combination of growth parameters.
STM Two versions of the VBGF[ were fitted. ] The first version was
t, L∞ is asymptotic FL, K is annual growth rate, and t0 is age
when Lt = 0. The usual VBGF was treated as a default growth
model. The{ second version was} the seasonally oscillating VBGF
Lt = L∞ 1 − e−[K (t −t0 )+St −St0 ] where St = 2CKπ sin 2π (t − ts )
and St0 = 2CKπ sin 2π (t0 − ts ). In the latter two expressions, the
parameter C is a constant indicating the amplitude of seasonal
growth oscillation, with 0 ≤ C ≤ 1 (C = 0 indicates constant
growth and C = 1 indicates seasonal cessation of growth). The
parameter ts , the so-called summer point, is the fraction of a
year (relative to the age of recruitment, t = 0) where the sine
wave oscillation begins (i.e., turns positive): ts typically starts the
seasonal cycle with respect to t = 0, and growth is considered
to be maximum (García and Duarte, 2006; Ba et al., 2016). The
seasonally oscillating VBGF was recommended by a reviewer
because the LFQ data appeared seasonally oscillating, and so was
devised to describe potential interannual variability in growth.
Note that, in the ELEFAN glossary, the notions of age and time are
used interchangeably. But ELEFAN methods shift the notion of t
from age to time so that, at recruitment for example, individuals
are assigned a length of zero and t0 becomes the time (i.e., date)
at that length rather than a theoretical age at length zero. The
same applies to ts as indicated previously.
ELEFAN algorithms operate on monthly LFQs but are designed
to analyze the progressions of LFQ modes over time assuming
that those modes represent age groups and describe sequential
growth of a cohort (i.e., a group of individuals born during the
same season in a year). Typically, the ages derived from modal
progressions using ELEFAN preclude the direct estimation of t0
and so are referred to as relative ages (Gayanilo et al., 1996).
To fit the growth model, ELEFAN applies numerous curves,
whereby the best combination of parameters produces a curve
that hits most modes, avoiding most troughs (see technical def-
inition below). This procedure involves the following four steps
(Isaac, 1990; Gayanilo and Pauly, 1997; Mildenberger et al., 2017):
(i) Restructuring the LFQ data by scoring the positive bins
(peaks), presumed to represent age groups, and the neg-
ative or zero bins (also called troughs) using the deviations
from a moving average across adjacent bins. Thus, peaks
are parts of the restructured LFQs that are above the run-
ning average; troughs are the opposite (in the ELEFAN
terminology, the words peak and mode are used inter-
changeably but peak appears to be mostly applicable to
restructured LFQs).
(ii) Calculating the sum of the highest values in positive peaks;
this sum is called available sum of peaks (ASP).
(iii) Calculating the estimated or explained sum of peaks (ESP),
which is the sum of all scores (positive and negative)
crossed by the estimated growth curves.
2
J. Munyandorero and D. Addis Regional Studies in Marine Science 39 (2020) 101468

Fig. 1. Areas around Florida, USA, that are continuously monitored with stratified-random sampling by the Fisheries Independent Monitoring program (FIM) of the
Fish and Wildlife Research Institute, Florida Fish and Wildlife Conservation Commission. Those areas are Apalachicola Bay (AP), Cedar Key (CK), Tampa Bay (TB),
Charlotte Harbor (CH), northern Indian River Lagoon (NIR), southern Indian River Lagoon (SIR), and northeast Florida (JX).

Table 2
Estimated parameters of the fork length (FL)-standard length (SL) relationships (FL = a + b SL) for combined sexes of marine forage fish of Florida, USA (the species
names were taken from www.fishbase.org; SL and FL are in cm).
Species Atlantic coast Gulf coast
n FL range a b r2 n FL range a b r2
Opisthonema oglinum 125 3.2–18.4 0.116 1.077 0.999 347 3.5–17.8 0.23 1.057 0.995
Anchoa mitchilli 759 1.9–7.4 0.086 1.066 0.989 837 1.9–7.5 0.028 1.085 0.993
Orthopristis chrysoptera 329 2–25 0.028 1.172 0.999 656 1.9–23.6 0.153 1.47 0.996
Lagodon rhomboides 1127 1.7–27.6 0.1 1.149 0.999 1388 1.5–28.8 0.106 1.146 0.998
Diplectrum formosum 115 3.2–19.8 0.139 1.168 0.99
Harengula jaguana 81 3.5–14.6 0.158 1.059 0.998 330 2.3–14.8 0.105 1.073 0.996
Mugil curema 427 2.5–30.7 0.151 1.153 0.999 350 2.8–45 0.343 1.139 0.998
Sardinella aurita 84 3.6–10.9 0.367 1.035 0.983
Leiostomus xanthurus 165 3.4–29 0.012 1.203 0.999 369 2.3–25 0.097 1.194 0.997
Eugerres plumieri 78 5.7–31.6 0.201 1.114 0.998 338 2–26.3 0.11 1.104 0.999
Mugil cephalus 1147 1.8–49 0.389 1.129 0.999 24253 2.2–57.5 1.706 1.068 0.965

Table 3
Estimated parameters of the total length (TL)-standard length (SL) relationships (TL = a + b SL) for combined sexes of marine forage fish of Florida, USA (the species
names were taken from www.fishbase.org; SL and TL are in cm).
Species Atlantic coast Gulf coast
n TL range a b r2 n TL range a b r2
Micropogonias undulates 383 1.5–42.2 0.535 1.202 0.998 555 1.7–34.1 0.449 1.207 0.998
Opisthonema oglinum 125 3.6–22.4 −0.377 1.344 0.999 349 3.9–21.7 −0.2 1.308 0.991
Anchoa mitchilli 794 2.1–8.2 0.023 1.203 0.991 985 1.9–8.9 −0.02 1.215 0.994
Orthopristis chrysoptera 358 2.2–27.2 −0.136 1.285 0.999 722 1.6–25.2 −0.015 1.26 0.996
Lagodon rhomboides 1153 1.6–32.4 −0.193 1.321 0.998 1592 1.5–33.1 −0.168 1.307 0.998
Diplectrum formosum 119 3.3–22 −0.199 1.327 0.983
Harengula jaguana 80 4–17.2 −0.007 1.261 0.997 332 2.5–17.6 −0.144 1.289 0.997
Mugil curema 441 2.7–34.4 0.035 1.292 0.997 356 2.9–49 0.046 1.289 0.997
Bairdiella chrysura 870 1.2–20.7 0,117 1.219 0.999 805 1.1–19.7 0.151 1.216 0.998
Sardinella aurita 83 3.9–12.4 −0.024 1.222 0.987
Leiostomus xanthurus 635 1.2–31.3 −0.153 1.29 0.999 2328 1.6–27 −0.08 1.277 0.997
Eugerres plumieri 77 7–37.7 0.538 1.355 0.996 337 2.2–31.8 0.097 1.327 0.997
Mugil cephalus 1178 2.1–54 0.193 1.272 0.998 24471 2.3–64.5 1.299 1.215 0.972

3
J. Munyandorero and D. Addis Regional Studies in Marine Science 39 (2020) 101468

Fig. 2. Restructured length–frequency (LFQ) data with superimposed curves of the usual von Bertalanffy growth function, obtained using the R package TropFishR
for forage fish on Florida (USA)’s Atlantic coast (see Table 1 for common and scientific names). Black bars in blue background and white bars in orange background
are, respectively, positive and negative deviations from moving averages. The x-axis reads as months from 2017 (17) but the monthly LFQs were aggregated across
1996–2017 .. (For interpretation of the references to color in this figure legend, the reader is referred to the web version of this article.)

Table 4
Estimated parameters of the fork length (FL)-standard length (TL) relationships (FL = a + b TL) for combined sexes of marine forage fish of Florida, USA (the species
names were taken from www.fishbase.org; FL and TL are in cm).
Species Atlantic coast Gulf coast
n FL range a b r2 n FL range a b r2
Opisthonema oglinum 125 3.2–18.4 0.425 0.8 0.999 350 3.5–17.8 0.457 0.804 0.995
Anchoa mitchilli 765 1.9–7.4 0.074 0.884 0.991 839 1.9–7.5 0.058 0.88 0.993
Orthopristis chrysoptera 330 2–25.2 0.146 0.913 0.999 653 1.8–23.6 0.168 0.911 0.998
Lagodon rhomboides 1133 1.5–27.6 0.3 0.868 0.999 1389 1.5–28.8 0.26 0.877 0.998
Diplectrum formosum 116 3.2–19.8 0.361 0.876 0.991
Harengula jaguana 80 3.5–14.6 0.166 0.839 0.995 330 2.3–14.8 0.27 0.829 0.997
Mugil curema 422 2.5–30.7 0.176 0.89 0.997 350 2.8–45 0.31 0.884 0.999
Sardinella aurita 84 3.6–10.94 0.407 0.845 0.99
Leiostomus xanthurus 164 3.4–29 0.138 0.928 0.999 368 2.3–25 −0.08 1.277 0.997
Eugerres plumieri 77 5.7–31.6 −0.341 0.827 0.996 340 2–26.3 0.048 0.831 0.998
Mugil cephalus 1147 2–49 0.201 0.888 0.999 24004 2.2–57.5 0.317 0.887 0.997

4
J. Munyandorero and D. Addis Regional Studies in Marine Science 39 (2020) 101468

Fig. 3. Restructured length–frequency (LFQ) data with superimposed curves of the usual von Bertalanffy growth function, obtained using the R package TropFishR
for forage fish on Florida (USA)’s Gulf coast (see Table 1 for common and scientific names). Black bars in blue background and white bars in orange background
are, respectively, positive and negative deviations from moving averages. The x-axis reads as months from 2017 (17) but the monthly LFQs were aggregated across
1996–2017 .. (For interpretation of the references to color in this figure legend, the reader is referred to the web version of this article.)

Table 5
Estimated parameters of the weight (W)–standard length (SL) relationships (W = aSLb ) for combined sexes of marine forage fish of Florida, USA (the species names
were taken from www.fishbase.org; W is in g and SL is in cm).
Species Atlantic coast Gulf coast
n SL range a (×10−3 ) b r2 n SL range a (×10−3 ) b r2
Micropogonias undulates 383 1.2–35.5 22 2.981 0.997 552 1.4–29 21 3.023 0.997
Opisthonema oglinum 122 2.9–17.3 23 2.95 0.997 346 3.1–16.7 16 3.096 0.988
Anchoa mitchilli 786 1.8–6.9 8 3.22 0.953 1285 1.6–7.1 7 3.323 0.969
Orthopristis chrysoptera 360 1.9–21.5 18 3.164 0.997 722 1.7–20.5 32 2.941 0.995
Lagodon rhomboides 1156 1.5–23.9 29 3.067 0.997 1608 1.2–25.2 27 3.076 0.996
Diplectrum formosum 118 2.7–16.8 20 3.101 0.992
Harengula jaguana 81 3.2–13.6 12 3.256 0.995 347 2.1–13.5 10 3.354 0.993
Mugil curema 464 2.1–26.8 21 3.006 0.996 358 2.3–40.2 24 2.956 0.996
Bairdiella chrysura 873 0.8–16.8 16 3.148 0.997 810 0.9–16.1 25 2.961 0.963
Sardinella aurita 84 3.2–10 7 3.331 0.996
Leiostomus xanthurus 634 1.2–24.3 15 3.239 0.997 2335 1.2–21 26 3.03 0.996
Eugerres plumieri 78 5–27.3 24 3.174 0.997 444 1.8–23.5 36 3.006 0.997
Mugil cephalus 1195 1.6–43.5 21 2.993 0.997

5
J. Munyandorero and D. Addis Regional Studies in Marine Science 39 (2020) 101468

Table 6
Estimated parameters of the weight (W)–fork length (FL) relationships (W = aFLb ) for combined sexes of marine forage fish of Florida, USA (the species names were
taken from www.fishbase.org; W is in g and FL is in cm).
Species Atlantic coast Gulf coast
n FL range a (×10−3 ) b r2 n FL range a (×10−3 ) b r2
Opisthonema oglinum 126 3.2–18.4 12 3.112 0.997 345 3.5–17.8 11 3.162 0.988
Anchoa mitchilli 757 2–7.4 6 3.241 0.956 833 1.9–7.5 6 3.256 0.973
Orthopristis chrysoptera 334 2–25 10 3.217 0.998 659 1.8–23.6 20 2.943 0.996
Lagodon rhomboides 1132 1.5–27.6 16 3121 0.998 1398 1.5–28.8 15 3.133 0.996
Diplectrum formosum 112 3.2–19.8 8 3.256 0.995
Harengula jaguana 80 3.5–14.6 12 3.158 0.996 330 2.3–14.8 8 3.331 0.993
Mugil curema 426 2.5–30.7 10 3.09 0.996 353 2.8–45 10 3.116 0.996
Sardinella aurita 84 3.6–10.9 5 3.403 0.949
Leiostomus xanthurus 166 3.4–26.9 8 3.233 0.995 366 2.3–25 11 3.165 0.996
Eugerres plumieri 78 5.7–31.6 19 3.133 0.997 337 2–26.3 25 3.02 0.998
Mugil cephalus 1152 1.8–49 10 3.095 0.997

(iv) Calculating an index of goodness-of-fit Rn = 10(ESP/ASP) /10
(0 < Rn ≤ 1). Rn is analogous to the correlation coefficient
in linear regressions. The greater the Rn, the more appro-
priate the combination of growth parameters for the LFQ
data.
ELEFAN methods fit LFQ data for generating the best com-
binations of L∞ and K only and, for the seasonally oscillating
VBGF, also the parameters C and ts . Instead of the VBGF param-
eter t0 , TropFishR returns a parameter called t_anchor, ranging
between and including 0 and 1. Typically, t_anchor is a time
point anchoring growth curves in year-length coordinate sys-
tem; it is considered to be reflecting a month of peak spawning
(e.g., t_anchor = 0.25 refers to April 1 of a year and would suggest
that spawning peaks in April). ELEFAN results are reliable if the
population samples are unbiased, which is herein likely because
the FIM program used stratified random surveys since 1996.
Unlike FISAT II, in which the moving average (MA) is fixed
at 5 size intervals regardless of length bin, TropFishR allows MA
to change depending on the LFQ data. In this case, a rule of
thumb for setting MA consists of using an approximate number
of bins spanning the smallest length–frequency width (i.e., the
youngest cohort) of the LFQ data (Taylor and Mildenberger, 2017).
Regardless, this study relied on the following criteria:
(i) A plus-length bin was created to include the fewer and
scattered counts of fish seen in some larger length classes.
This precluded any artifact of higher relative ages resulting
from peaks that otherwise might occur at those extreme
lengths.
(ii) MA was a value such that the growth curves would (a) span
most positive length bins and (b) produce the highest Rn
value. Furthermore, given the maximum lengths in samples
(Lmax ) and consistent with the questionability of L∞ in
FishBase (www.fishbase.org), L∞ would fall within Lmax ±
1/3Lmax .
In running ELEFAN_GA, the default, starting value of MA was
5. However, the growth parameters derived using ELEFAN_GA
are very sensitive to the choice of MA. In fact, to determine
the most preferable MA value, the more MA is increased, the
more the restructured LFQs with positive peaks are truncated
and associated with fewer growth curves, and the lower is L∞
(eventually less than Lmax − 1/3Lmax ). The opposite occurs when
MA is reduced; this can be associated with C > 1, indicating the
so-called periods of shrinkage in length (Taylor and Mildenberger,
2017) and, apparently whatever the VBGF version, with unreliable
lower K and higher L∞ and Amax estimates. A value of MA leading
to the shrinkage in length was rejected; an estimate of L∞ < Lmax
− 1/3Lmax following a reduced MA value was deemed unreliable.
6
Given the parameters of L∞ and K, natural mortality M was
estimated with TropFishR using Pauly’s nonlinear empirical equa-
tion as updated by Then et al. (2015): M = 4.118K 0.73 L− ∞
0.33
.
The estimated values of growth parameters and natural mortality
were plotted against each other to verify their conformity with
the patterns of life-history expectations (e.g., Adams, 1980).
3. Results
3.1. Morphometric relationships
Length–length and weight–length relationships were devel-
oped for 12 forage fish species on the Atlantic coast of Florida
and for 13 forage fish species on the Gulf coast of Florida (Ta-
bles 2–5). All relationships were significant (p < 0.01) and as-
sociated with high coefficients of determination (r 2 ). For some
species (e.g., Atlantic croaker, Micropogonias undulates, and Silver
perch, Bairdiella chrysura), the development of some morpho-
metric relationships were not possible because of the lack of
data.
3.2. Growth parameters
The growth curves for some forage fish of Florida are super-
imposed on restructured LFQ data, characterized by black bars
(along with a blue background) and white bars (along with an
orange background) for, respectively, positive and negative devia-
tions from the moving average (Figs. 2–5). The best combinations
of the VBGF parameters are in Tables 8 and 9 along with the
moving average used, the goodness-of-fit parameter Rn, t-anchor,
and other growth traits. L∞ values based on both the usual and
seasonally oscillating VBGFs for bay anchovy (Anchoa mitchilli)
on the Atlantic coast and Atlantic thread herring (Opisthonema
oglinum) on the Gulf coast may be unreliable because they are
less than Lmax − 1/3Lmax . Following the latter criterion, the fit
of the seasonally oscillating VBGF to LFQ data of scaled sardine
(Harengula jaguana) on the Gulf coast also produced an unreliable
L∞ value.
Overall, the parameter Rn is similar for all VBGF fits, but Rn
from the seasonally oscillating VBGF increased for most species,
thereby suggesting best fits to LFQ data (i.e., that growth function
tracked better the progressions of modes). However, there is little
difference between estimates of L∞ , Amax , and occasionally K from
the two VBGF versions (Tables 8 and 9). The values of t_anchor
suggest that the time at which length equals zero is in the second
half of the year for most forage fish on both coasts of Florida.
Through the parameter C of the seasonally oscillating VBGF, the
strength of growth oscillation (i.e., the deviation from the usual
VBGF fit) is noticeable (C = 0.31–0.87), except for sand perch
(Diplectrum formosum) on the Gulf coast for which C = 0.07.
Given that the parameter ts typically reflects the time of year
J. Munyandorero and D. Addis Regional Studies in Marine Science 39 (2020) 101468

Fig. 4. Restructured length–frequency (LFQ) data with superimposed curves of the seasonally oscillating von Bertalanffy growth function, obtained using the R
package TropFishR for forage fish on Florida (USA)’s Atlantic coast (see Table 1 for common and scientific names). Black bars in blue background and white bars in
orange background are, respectively, positive and negative deviations from moving averages. The x-axis reads as months from 2017 (17) but the monthly LFQs were
aggregated across 1996–2017 .. (For interpretation of the references to color in this figure legend, the reader is referred to the web version of this article.)

Table 7
Estimated parameters of the weight (W)–total length (TL) relationships (W = aTLb ) for combined sexes of marine forage fish of Florida, USA (the species names
were taken from www.fishbase.org; W is in g and TL is in cm).
Species Atlantic coast Gulf coast
n TL range a (×10−3 ) b r2 n TL range a (×10−3 ) b r2
Micropogonias undulates 379 1.7–42.2 8 3.119 0.997 552 1.7–34.1 6 3.232 0.996
Opisthonema oglinum 125 3.6–22.4 9 2.994 0.997 346 3.9–21.7 10 2.996 0.989
Anchoa mitchilli 791 2.1–8.2 4 3.237 0.953 983 1.9–8.9 5 3.232 0.975
Orthopristis chrysoptera 360 2.3–27.2 8 3.208 0.998 717 2–25.2 17 2.916 0.996
Lagodon rhomboides 1151 1.6–32.4 15 3.027 0.998 1600 1.5–31.3 13 3.068 0.997
Diplectrum formosum 118 3.3–22 5 3.257 0.992
Harengula jaguana 81 4–17.2 8 3.12 0.995 330 2.5–17.6 5 3.297 0.996
Mugil curema 443 2.7–34.4 8 3.078 0.996 356 2.9–49 5 3.189 0.996
Bairdiella chrysura 874 0.9–20.7 7 3.218 0.997 806 1.2–19.7 10 3.068 0.997
Sardinella aurita 84 3.9–12.4 5 3.164 0.963
Leiostomus xanthurus 629 1.5–31.3 8 3.187 0.997 2314 1.6–27 11 3.065 0.996
Eugerres plumieri 77 7–37.7 6 3.296 0.997 337 2.2–31.8 14 3.023 0.997
Mugil cephalus 1186 2.1–54 6 3.129 0.997

7
J. Munyandorero and D. Addis Regional Studies in Marine Science 39 (2020) 101468

Fig. 5. Restructured length–frequency (LFQ) data with superimposed curves of the seasonally oscillating von Bertalanffy growth function, obtained using the R
package TropFishR for forage fish on Florida (USA)’s Gulf coast (see Table 1 for common and scientific names). Black bars in blue background and white bars in
orange background are, respectively, positive and negative deviations from moving averages. The x-axis reads as months from 2017 (17) but the monthly LFQs were
aggregated across 1996–2017 .. (For interpretation of the references to color in this figure legend, the reader is referred to the web version of this article.)

when growth turns positive (Taylor and Mildenberger, 2017, e.g.,
ts = 0.5 means that growth turns positive on 1st July), then,
except for bay anchovy and scaled sardine (for which L∞ is
unreliable), sine wave oscillations may begin in one of the spring
and summer months depending on the species (ts = 0.32–63). Fi-
nally, across-species growth rates (K ) are, as expected, negatively
correlated with L∞ ; likewise, across-species maximum lengths
(Lmax ) and maximum relative ages (Amax ) are positively related
to L∞ (Fig. 6(a–f)).
3.3. Natural mortality
Estimates of natural mortality are smaller for larger (higher
Lmax ) and older (higher Amax ) forage fish (Tables 8 and 9, Fig. 6(g)
and (h)).
8
4. Discussion
The primary objective of this study was to develop the mor-
phometric relationships, growth parameters, and natural mortal-
ity rates for forage fish species of Florida (USA). Such a study may
raise interest in the opportunity offered by the availability of FIM
length data for obtaining non-parametric life-history parameters
of so many of Florida’s inshore fish species for which minimal
biological samples, essentially in terms of size measurements,
have been collected. In the end, the life-history variables thus
generated could be useful when species-specific stock assess-
ments or evaluations of ecosystem management strategies are
contemplated for Florida marine waters.
The morphometric equations were intended to generate stand-
alone parameters for forage fish of Florida, but the estimated
parameters and the underlying size ranges and maximum sizes
were consistent with parameter values in FishBase (www.fish
base.org), except for Spanish sardine (Sardinella aurita), striped
mojarra (Eugerres plumieri), and spot croaker (Leiostomus xanthu-
rus). For Spanish sardine, the maximum total lengths and size
J. Munyandorero and D. Addis Regional Studies in Marine Science 39 (2020) 101468

Fig. 6. Scatterplots of the relationships between (a and b) growth rate K, (c and d) maximum relative age Amax , (e and f) maximum length Lmax , and (g and h) natural
mortality M against asymptotic length L∞ estimated using the usual von Bertalanffy growth function (left panels) and the seasonally oscillating von Bertalanffy
growth function (right panels) for forage fish of Florida (USA)’s Atlantic coast (circles) and Gulf coast (triangles).

ranges were far smaller than the values reported in FishBase
and Ohs et al. (2013), suggesting that the FIM program may not
be targeting adult individuals, which usually are pelagic, and
that the estimated parameters relate to juveniles. For striped
mojarra, this study reports a maximum total length larger than
those available in FishBase; for spot croaker, there were limited
parameters in FishBase for comparison. Regarding the weight–
length relationships, most exponents in this study were close
to or exceeded 3, indicating that forage fish in Florida have an
isometric or a positive-allometric growth pattern.
The usual VBGF was a priori treated as a default growth model
to fit the LFQ data of forage fish in Florida because, in most appli-
cations, the parameters of such a growth function have commonly
been the basis for calculating mean weight at age. The LFQs in
question also showed temporal progressions of modes suggesting
seasonally oscillating VBGF curves for those fish. Such growth
patterns and functions have been assumed and used elsewhere
(e.g., Moreau and Nyakageni, 1992; Pauly et al., 1995; Acosta,
2000; García and Duarte, 2006; Nwosu et al., 2010; Ba et al.,
2016), and have been attributed to changes in environmental
factors or in food availability (Morales-Nin and Panfili, 2005;
9
Vahabnezhad et al., 2020). Alternatively, some LFQ data suggested
two possible cohorts (i.e., two groups of fish born during two
different seasons in a year, each with a unique growth curve)
as in Moreau et al. (1991). The fittings of the seasonally oscil-
lating VBGFs were performed only because the progression of
modes suggested seasonal variations of growth, but any support-
ing causal environmental and biological factors remain unknown.
Finally, although ELEFAN in TropFishR can eventually fit growth
curves for multiple cohorts, such cohort-specific growth fits were
not attempted because there was no evidence that the species
studied have spawning dynamics resulting in more than one
cohort per year.
Using length in determining age and growth of marine or-
ganisms are usually based on subjective criteria and unjusti-
fied assumptions (Pauly and Morgan, 1987; Isaac, 1990; Gul-
land and Rosenberg, 1992; Taylor and Mildenberger, 2017). A
question, therefore, is whether the ELEFAN-estimated relative
ages and growth parameters for Florida’s forage fish are reli-
able. Although, due to space limitations, the maximum observed
ages obtained using other aging techniques are not reported, the
ELEFAN method may overestimate maximum age, especially for
J. Munyandorero and D. Addis Regional Studies in Marine Science 39 (2020) 101468

Table 8
Estimated parameters of the usual von Bertalanffy growth function for marine forage fish of Florida, USA. The table also includes the
moving average (MA) used to estimate those parameters as well as the maximum length observed in LFQs (Lmax ) and the maximum
relative age estimated by TropFishR (Amax ). L∞ and Lmax are in cm; Amax is in years; and K and M are per year. L∞ for species
marked by an asterisk (*) is less than Lmax – 1/3Lmax and, hence, is unreliable according to www.fishbase.org.
Species MA L∞ K t_anchor Rn M Lmax Amax
Atlantic coast
Micropogonias undulates 3 48.7 0.28 0.66 0.24 0.45 52.0 11
Opisthonema oglinum 3 25.1 0.30 0.80 0.31 0.6 26.8 10
Anchoa mitchilli* 3 6.0 0.63 0.68 0.27 1.63 10.1 4
Orthopristis chrysoptera 5 34.3 0.17 0.75 0.35 0.36 40.9 18
Lagodon rhomboides 3 29.5 0.28 0.83 0.36 0.53 35.1 11
Harengula jaguana 3 15.6 0.42 0.71 0.27 0.87 21.5 8
Mugil curema 3 47.3 0.33 0.22 0.22 0.52 48.0 10
Bairdiella chrysura 3 26.9 0.45 0.56 0.20 0.78 32.2 7
Leiostomus xanthurus 5 33.2 0.4 0.91 0.27 0.66 37.0 8
Eugerres plumieri 5 29.3 0.45 0.24 0.22 0.75 34.8 7
Mugil cephalus 3 57.4 0.14 0.69 0.23 0.26 64.0 21
Gulf coast
Micropogonias undulates 3 40.3 0.40 0.79 0.29 0.63 48.0 8
Opisthonema oglinum* 7 17.2 0.53 0.05 0.41 1.02 27.8 4
Anchoa mitchilli 3 10.2 0.59 0.44 0.27 1.3 13.6 6
Orthopristis chrysoptera 3 29.5 0.3 0.77 0.33 0.55 40.3 11
Lagodon rhomboides 3 28.9 0.3 0.84 0.37 0.56 32.5 10
Diplectrum formosum 5 18.7 0.49 0.1 0.22 0.93 23.5 7
Harengula jaguana 3 19.0 0.51 0.12 0.28 0.96 25.2 6
Mugil curema 3 45.3 0.46 0.22 0.31 0.66 42.0 7
Bairdiella chrysura 3 32.3 0.31 0.70 0.27 0.56 36.2 10
Leiostomus xanthurus 3 33.1 0.23 0.71 0.28 0.45 36.0 13
Eugerres plumieri 3 32.3 0.33 0.22 0.24 0.58 36.7 10
Mugil cephalus 3 55.3 0.19 0.71 0.22 0.33 57.5 16

Table 9
Estimated parameters of the seasonally oscillating von Bertalanffy growth function for marine forage fish of Florida, USA. The moving
average (MA) and the maximum length observed in LFQs (Lmax ) were the same as in Table 8. The table also includes the maximum
relative age estimated by TropFishR (Amax ). Amax is in years and K and M are per year. L∞ for species marked by an asterisk (*) is
less than Lmax – 1/3Lmax and, hence, is unreliable according to www.fishbase.org.
Species L∞ K t_anchor C ts Rn M Amax
Atlantic coast
Micropogonias undulates 51.6 0.26 0.62 0.83 0.41 0.32 0.42 12
Opisthonema oglinum 24.4 0.32 0.71 0.31 0.47 0.29 0.63 10
Anchoa mitchilli* 5.9 0.66 0.61 0.67 0.15 0.29 1.68 5
Orthopristis chrysoptera 32.8 0.18 0.65 0.41 0.50 0.34 0.37 17
Lagodon rhomboides 29.4 0.35 0.82 0.45 0.56 0.38 0.63 9
Harengula jaguana 16.0 0.45 0.70 0.49 0.34 0.25 0.92 7
Mugil curema 48.6 0.3 0.81 0.87 0.55 0.30 0.47 11
Bairdiella chrysura 27.6 0.37 0.32 0.39 0.55 0.22 0.66 9
Leiostomus xanthurus 32.7 0.40 0.73 0.66 0.44 0.36 0.66 8
Eugerres plumieri 33.2 0.57 0.69 0.56 0.35 0.20 0.87 6
Mugil cephalus 57.3 0.15 0.6 0.84 0.51 0.33 0.27 21
Gulf coast
Micropogonias undulates 44.3 0.39 0.68 0.66 0.41 0.4 0.59 8
Opisthonema oglinum* 17.5 0.61 0.46 0.83 0.51 0.48 1.12 5
Anchoa mitchilli 10.8 0.35 0.40 0.66 0.63 0.31 0.88 9
Orthopristis chrysoptera 28.6 0.46 0.75 0.65 0.49 0.42 0.78 7
Lagodon rhomboides 28.6 0.3 0.67 0.59 0.5 0.42 0.57 11
Diplectrum formosum 17.9 0.81 0.46 0.07 0.35 0.23 1.34 4
Harengula jaguana* 14.0 0.79 0.18 0.71 0.27 0.63 1.46 4
Mugil curema 42.0 0.53 0.22 0.51 0.59 0.26 0.75 6
Bairdiella chrysura 33.7 0.24 0.8 0.54 0.56 0.34 0.46 13
Leiostomus xanthurus 32.6 0.24 0.69 0.46 0.36 0.31 0.47 13
Eugerres plumieri 32.2 0.35 0.35 0.63 0.32 0.24 0.61 9
Mugil cephalus 56.5 0.15 0.65 0.57 0.58 0.27 0.27 20

short-lived species. For example, otolith reading indicated (i) that
Atlantic thread herring of age-3+ in the Chesapeake Bay, USA
(Newberger and Houde, 1995) and of age-1+ in Florida waters
(Pierce et al., 2001) were rare and (ii) that pinfish in Tampa
Bay, Florida, reached a maximum age of 7 years (Nelson, 2002).
Conversely, ELEFAN-based estimates of growth parameters are
generally comparable with results obtained using hard parts such
as otoliths and scales (Supplementary data in Table S1). Historical
growth parameters in Table S1 are probably not exhaustive; they
10
could not particularly be documented for pigfish and silver perch
of any location.
5. Conclusion
The present study relied on the size data collected by the
Fishery Independent Monitoring program of the Florida Fish and
Wildlife Conservation Commission’s Fish and Wildlife Research
Institute to develop Florida-specific morphometric relationships
J. Munyandorero and D. Addis
and length-based growth parameters and curves of Florida forage
fish. Overall, the ages and growth parameters estimated herein
should be considered preliminary but still useful, pending possi-
ble improvement from other information such as tagged individ-
uals or the reading of hard parts.
CRediT authorship contribution statement
Joseph Munyandorero: Formal analysis, Writing - original
draft. Dustin Addis: Formal analysis, Writing - original draft.
Declaration of competing interest
The authors declare that they have no known competing finan-
cial interests or personal relationships that could have appeared
to influence the work reported in this paper.
Acknowledgments
A. Acosta and C. Bradshaw provided comments on the early
version of this paper. FIM scientists are specially thanked for
having tirelessly conducted surveys and collected data around
Florida. R. Grunwald and B. Crowder helped in the preparation of
this paper. This study was prepared under award
NA17NMF4740329 from the National Marine Fisheries Service
of the National Oceanic and Atmospheric Administration to the
Florida Fish and Wildlife Conservation Commission.
Appendix A. Supplementary information
Supplementary material related to this article can be found
online at https://doi.org/10.1016/j.rsma.2020.101468.
References
Acosta, A., 2000. Estimation of growth and mortality of bay anchovy, Anchoa
mitchilli. In: Florida Bay, Florida, U.S.A. Proc. 51st Gulf Caribb. Fish. Inst. vol.
51, pp. 204–214.
Adams, P.B., 1980. Life history patterns in marine fishes and their consequences
for fisheries management. Fish. Bull. 78, 1–12.
Adams, G.D., Leaf, R.T., Ballenger, J.C., Arnott, S.A., McDonough, C.J., 2018. Spatial
variability in the growth of sheepshead (Archosargus probatocephalus) in the
southeast US: implications for assessments and management. Fish. Res. 206,
35–43.
Ba, K., Thiaw, M., Lazar, N., Sarr, A., Brocher, T., Ndiaye, I., Faye, A., Sadio, O.,
Panfili, J., Thiaw, O.T., Brehmer, P., 2016. Resilience of key biological parame-
ters of the Senegalese flat sardinella to overfishing and climate change. PLoS
One 11 (6), http://dx.doi.org/10.1371/journal.pone.0156143.
Bortone, S.A., 2003. Biology of the Spotted Seatrout. CRC Press, Boca Raton.
Brown-Peterson, N.J., Burns, K.M., Overstreet, R.M., 2009. Regional differences in
Florida red snapper reproduction. In: Proc. 61st Gulf Caribb. Fish. Inst., vol.
61, pp. 149–155.
Crabtree, R.E., Bullock, L.H., 1998. Age, growth, and reproduction of black
grouper, Mycteroperca bonaci, in Florida. Fish. Bull. 96, 735–753.
García, C.B., Duarte, L.O., 2006. Length–based estimates of growth parameters
and mortality rates of fish populations of the Caribbean Sea. J. Appl. Ichthyol.
22, 193–200.
Gayanilo, F.C., Pauly, D., 1997. FAO-ICLARM Stock Assessment Tools. Reference
Manual. In: FAO Computerized Information Series (Fisheries), vol. 8, FAO,
Rome.
Gayanilo, F.C., Sparre, P., Pauly, D., 1996. FAO-ICLARM Stock Assessment Tools
(FiSAT) User’s Manual. In: FAO Computerized Information Series (Fisheries),
vol. 8, FAO, Rome.
11
Regional Studies in Marine Science 39 (2020) 101468
Grüss, A., Harford, W.J., Schirripa, M.J., Velez, L., Sagarese, S.R., Shin, Y.-J.,
Verley, P., 2016. Management strategy evaluation using the individual-based,
multispecies modeling approach OSMOSE. Ecol. Model. 340, 86–105.
Gulland, .J.A., Rosenberg, A.A., 1992. A Review of Length-Based Approaches to
Assessing Fish Stocks. In: FAO Fisheries Technical Paper 323, FAO, Rome.
Herdter, E., Peebles, E., Mahmoudi, B., Murawski, S., 2019. Spatial variability in
size-structure, growth, and recruitment of spotted seatrout among six florida
estuaries. Mar. Coast. Fish. 11, 97–111.
Hilborn, R., Amoroso, R.O., Bogazzi, E., Jensen, O.P., Parma, A.M., Szuwalski, C.,
Walters, C.J., 2017. When does fishing forage species affect their predators?.
Fish. Res. 191, 211–221.
Isaac, V.J., 1990. The Accuracy of Some Length-Based Methods for Fish Population
Studies. ICLARM Technical Report 27, ICLARM, Manila.
Koehn, L.E., Essington, T.E., Marshall, K.N., Sydeman, W.J., Szoboszlai, A.I.,
Thayer, J.A., 2017. Trade-offs between forage fish fisheries and their
predators in the California Current. ICES J. Mar. Sci. 74, 2448–2458.
Mildenberger, T.K., Taylor, M.H., Wolff, M., 2017. TropFishR: An R package for
fisheries analysis with length-frequency data. Meth. Ecol. Evol. 8, 1520–1527.
Morales-Nin, B., Panfili, J., 2005. Seasonality in the deep sea and tropics revisited:
What can otoliths tell us? Mar. Freshw. Res. 56, 585–598.
Moreau, J., Munyandorero, J., Nyakageni, B., 1991. Evaluation des paramètres
démographiques chez Stolothrissa tanganykae et Limnothrissa miodon du lac
Tanganyika. Verh. int. Ver. Limnol. 24, 2552–2558.
Moreau, J., Nyakageni, B., 1992. Luciolates stappersi in Lake Tanganyika. Demo-
graphical status and possible recent variations assessed by length frequency
distributions. Hydrobiologia 232, 57–64.
Nelson, G.A., 2002. Age, growth, mortality and distribution of pinfish (Lagodon
rhomboides) in Tampa Bay and adjacent Gulf of Mexico waters. Fish. Bull.
100, 582–592.
Newberger, T.A., Houde, E.D., 1995. Population biology of bay anchovy Anchoa
mitchilli in the mid Chesapeake Bay. Mar. Ecol. Prog. Ser. 116, 25–37.
Nwosu, F.M., Enin, U.I., Holzlöhner, S., 2010. Assessment of artisanal fishery
impacts on West African croaker Pseudotolithus elongatus in the Cross River
Estuary, Nigeria, using length-based models. N. Am. J. Fish. Manage. 30,
860–865.
Ohs, C.L., Creswell, R.L., Dimaggio, M.A., 2013. Growing Marine Baitfish. a Guide
of Florida’s Common Marine Baitfish and their Potential for Aquaculture.
University of Florida, Fort Pearce. Sea Grant Florida.
Pauly, D., Moreau, J., Abad, N., 1995. Comparison of age-structured and length-
converted catch curves of brown trout Salmo trutta in two French rivers.
Fish. Res. 22, 197–204.
Pauly, D., Morgan, G.R., 1987. Length-based methods in Fisheries research. In:
ICLARM. Conference Proceedings, vol. 13. ICLARM, Manila.
Pierce, D.J., Mahmoudi, B., Wilson, R.R., 2001. Age and growth of the scaled her-
ring, harengula jaguana, from Florida waters, as indicated by microstructure
of the sagittae. Fish. Bull. 99, 202–209.
Taylor, M.H., Mildenberger, T.K., 2017. Extending electronic length frequency
analysis in R. Fish. Manage. Ecol. 24, 330–338.
Then, A.Y., Hoenig, J.M., Hall, N.G., Hewitt, D.A., 2015. Evaluating the predic-
tive performance of empirical estimators of natural mortality rate using
information on over 200 fish species. ICES J. Mar. Sci. 72, 82–92.
Vahabnezhad, A., Taghavimotlagh, S.A., Salarpouri, A., 2020. Estimation of growth
parameters and mortality rate of Upeneus sulphureus (Cuvier, 1829) in the
Persian Gulf ecosystem. J. Surv. Fish. Sci. 7, 69–81.
Walters, C., Martell, S.J.D., Christensen, V., Mahmoudi, B., 2008. An ecosim model
for exploring gulf of Mexico ecosystem management options: Implications
of including multistanza life-history models for policy predictions. Bull. Mar.
Sci. 83, 251–271.
Wang, K., Zhang, C., Xu, B., Xiu, W., Re, Y., 2020. Selecting optimal bin size
to account for growth variability in Electronic LEngth Frequency ANalysis
(ELEFAN). Fish. Res. 225, http://dx.doi.org/10.1016/j.fishres.2019.105474.
White, D.B., Palmer, S.M., 2004. Age, growth, and reproduction of the red
snapper, Lutjanus campechanus, from the Atlantic waters of the southeastern
U.S. Bull. Mar. Sci. 75, 335–360.
Young, J.M., Yeiser, B.G., Whitthington, J.A., Dutka-Gianelli, J., 2020. Maturation of
female common snook centropomus undecimalis: implications for managing
protandrous fishes. J. Fish Biol. (in press). http://dx.doi.org/10.1111/jfb.14475.