Short Communication
Short Communication
Length–weight relations of 39 continental-shelf and deep-water fishes (Actinopterygii) from northwestern Gulf of México
expand article infoAriel A. Chi-Espínola, M. Eugenia Vega-Cendejas, J. Mirella Hernández De Santillana
‡ Laboratorio de Taxonomía y Ecología de Peces, Mérida, Mexico
Open Access


Fishes from the northwestern Gulf of Mexico were surveyed during four oceanographic campaigns (February and October 2016, June and September 2017) using a shrimp trawl net and benthic sled net in 20 locations at depths that ranged from 43 to 3608 m. Length–weight relations (LWR) were estimated for 39 fish species (in alphabetical order): Bembrops gobioides (Goode, 1880); Centropristis philadelphica (Linnaeus, 1758); Chauliodus sloani Bloch et Schneider, 1801; Chlorophthalmus agassizi Bonaparte, 1840; Chloroscombrus chrysurus (Linnaeus, 1766); Citharichthys spilopterus Günther, 1862; Coelorinchus caelorhincus (Risso, 1810); Cyclopsetta chittendeni Bean, 1895; Cyclothone alba Brauer, 1906; Cyclothone braueri Jespersen et Tåning, 1926; Cyclothone pseudopallida Mukhacheva, 1964; Dibranchus atlanticus Peters, 1876; Epigonus pandionis (Goode et Bean, 1881); Fowlerichthys radiosus (Garman, 1896); Laemonema goodebeanorum Meléndez et Markle, 1997; Lagocephalus laevigatus (Linnaeus, 1766); Lepophidium brevibarbe (Cuvier, 1829); Lutjanus campechanus (Poey, 1860); Malacocephalus occidentalis Goode et Bean, 1885; Merluccius albidus (Mitchill, 1818); Micropogonias furnieri (Desmarest, 1823); Monolene sessilicauda Goode, 1880; Ogcocephalus declivirostris Bradbury, 1980; Peristedion greyae Miller, 1967; Porichthys plectrodon Jordan et Gilbert, 1882; Prionotus longispinosus Teague, 1951; Prionotus paralatus Ginsburg, 1950; Pristipomoides aquilonaris (Goode et Bean, 1896); Rhynchoconger flavus (Goode et Bean, 1896); Sardinella aurita Valenciennes, 1847; Saurida brasiliensis Norman, 1935; Sternoptyx diaphana Hermann, 1781; Symphurus diomedeanus (Goode et Bean, 1885); Synagrops bellus (Goode et Bean, 1896); Trachurus lathami Nichols, 1920; Trichiurus lepturus Linnaeus, 1758; Trichopsetta ventralis (Goode et Bean, 1885); Urophycis cirrata (Goode et Bean, 1896); Zalieutes mcgintyi (Fowler, 1952). The fish species studied represented 28 families (in alphabetical order): Antennariidae, Batrachoididae, Bembropidae, Bothidae, Carangidae, Chlorophthalmidae, Congridae, Cyclopsettidae, Cynoglossidae, Dorosomatidae, Epigonidae, Gonostomatidae, Lutjanidae, Macrouridae, Merlucciidae, Moridae, Ogcocephalidae, Ophidiidae, Phycidae, Sciaenidae, Serranidae, Sternoptychidae, Stomiidae, Synagropidae, Synodontidae, Tetraodontidae Trichiuridae, Triglidae. A new maximum standard length (SL) was recorded for Cyclothone alba, C. braueri, C. pseudopallida, and Lepophidium brevibarbe. A positive allometric growth was reported in nine species, negative allometric growth in 16 species, and isometric growth in 14 species.


bathyal, continental shelf, deep-water fish, Gulf of Mexico, length–weight relation


Currently, demersal fishes in the northwestern Gulf of Mexico are under pressure from a growing industry focusing on oil exploration and extraction (Patiño-Ruiz et al. 2003). They are also affected by trawling, forming part of the discarded fauna from shrimp fishing in the area (Chávez-López and Morán-Silva 2019). One way to assess the scope and impact of these activities on biodiversity is by drawing up a list of the fish fauna in the area, as well as determining the affected life cycles, which are identified by studying the sizes of the fish specimens (Hernández-Padilla et al. 2020). For this process, length–weight relation (LWR) analyses are used, which commonly focus on identifying fish stocks, and the growth rate of a particular species, among others (Sandoval-Huerta et al. 2015). Therefore, the presently reported study was intended to determine the LWR of 39 dominant fish species from the northwestern region of the Gulf of Mexico in areas ranging from the continental shelf to the bathyal zone.

Materials and methods

Data collection was carried out during four oceanographic study surveys aboard the research vessel RV JUSTO SIERRA, each trip with an approximate duration of 10 days during the months of February and October 2016, and June and September 2017 (adequate weather conditions and project logistics). The activity was carried out at 20 sampling sites comprising depths between 43 and 3608 m. Two types of fishing gear were implemented, depending on the depth of each site, a shrimp trawl (18.29 m long and 4.57 cm mesh size) for depths between 50 and 500 m (9 sites) and a benthic sled net (32.4 m long and 2.5 cm mesh size) for depths between 500 and 3608 m (11 sites); both nets were hauled for one mile at a constant speed of 2.7 knots.

The collected fishes were labeled and immediately frozen at –20°C. They were subsequently transferred to the laboratory, where they were identified using specialized references (Carpenter 2002a, 2002b; McEachran and Fechhelm 2005). Individual weight and standard length (SL) were determined for all specimens and supplemented with the relevant site information, such as the coordinates, and depth. All specimens were measured and weighed fresh, fixed, and preserved in 80% ethyl alcohol. Some fish individuals were deposited in the ichthyological collection (CINV-NEC) of CINVESTAV-Merida in Mexico. The following 39 species, representing 28 families were investigated (Table 1), including Rhynchoconger flavus (Goode et Bean, 1896) [Congridae]; Sardinella aurita Valenciennes, 1847 [Dorosomatidae]; Cyclothone alba Brauer, 1906, Cyclothone braueri Jespersen et Tåning, 1926, Cyclothone pseudopallida Mukhacheva, 1964 [Gonostomatidae]; Sternoptyx diaphana Hermann, 1781 [Sternoptychidae]; Chauliodus sloani Bloch et Schneider, 1801 [Stomiidae]; Saurida brasiliensis Norman, 1935 [Synodontidae]; Chlorophthalmus agassizi Bonaparte, 1840 [Chlorophthalmidae]; Coelorinchus caelorhincus (Risso, 1810), Malacocephalus occidentalis Goode et Bean, 1885 [Macrouridae]; Laemonema goodebeanorum Meléndez et Markle, 1997 [Moridae]; Merluccius albidus (Mitchill, 1818) [Merlucciidae]; Urophycis cirrata (Goode et Bean, 1896) [Phycidae]; Lepophidium brevibarbe (Cuvier, 1829) [Ophidiidae]; Porichthys plectrodon Jordan et Gilbert, 1882 [Batrachoididae]; Chloroscombrus chrysurus (Linnaeus, 1766), Trachurus lathami Nichols, 1920 [Carangidae]; Citharichthys spilopterus Günther, 1862, Cyclopsetta chittendeni Bean, 1895 [Cyclopsettidae]; Monolene sessilicauda Goode, 1880, Trichopsetta ventralis (Goode et Bean, 1885) [Bothidae]; Symphurus diomedeanus (Goode et Bean, 1885) [Cynoglossidae]; Trichiurus lepturus Linnaeus, 1758 [Trichiuridae]; Bembrops gobioides (Goode, 1880) [Bembropidae]; Synagrops bellus (Goode et Bean, 1896) [Synagropidae]; Epigonus pandionis (Goode et Bean, 1881) [Epigonidae]; Centropristis philadelphica (Linnaeus, 1758) [Serranidae]; Lutjanus campechanus (Poey, 1860), Pristipomoides aquilonaris (Goode et Bean, 1896) [Lutjanidae]; Prionotus longispinosus Teague, 1951, Prionotus paralatus Ginsburg, 1950, Peristedion greyae Miller, 1967 [Triglidae]; Micropogonias furnieri (Desmarest, 1823) [Sciaenidae]; Fowlerichthys radiosus (Garman, 1896) [Antennariidae]; Dibranchus atlanticus Peters, 1876, Ogcocephalus declivirostris Bradbury, 1980, Zalieutes mcgintyi (Fowler, 1952) [Ogcocephalidae]; and Lagocephalus laevigatus (Linnaeus, 1766) [Tetraodontidae].

Table 1.

Length–weight relations for 39 fish species caught in northwestern Gulf of México.

Species Depth [m] n SL [cm] Weight [g] a 95% CI a b 95% CI b Growt type R 2 Reference data
L m [cm] L max [cm]
Rhynchoconger flavus 35 14.2–42.7 4.4–133.0 0.0012 0.001–0.003 3.055 2.817–3.293 I 0.954 150.0TL
Sardinella aurita 51 7.0–19.3 4.1–99.3 0.0124 0.007–0.022 3.024 2.831–3.216 I 0.953 12.0TL 41.0TL
Cyclothone alba ≥500 75 1.3–5.6 0.02–0.42 0.0076 0.007–0.009 2.309 2.168–2.449 –A 0.936 1.56SL2 2.9SL
Cyclothone braueri ≥500 22 1.4–4.6 0.02–0.23 0.0045 0.002–0.005 3.000 I 0.975 2.0SL,2 3.8SL
Cyclothone pseudopallida ≥500 71 1.5–4.8 0.02–0.51 0.0076 0.006–0.009 2.518 2.333–2.703 –A 0.914 1.75SL,2 4.6SL
Sternoptyx diaphana ≥500 26 1.2–4.5 0.09–4.21 0.0503 0.041–0.062 2.892 2.671–3.114 I 0.968 5.5SL
Chauliodus sloani ≥500 25 4.5–19.2 0.09–17.03 0.0012 0.001–0.002 3.181 2.919–3.442 +A 0.965 15.1SL,3 35.0SL
Saurida brasiliensis 203 3.1–9.7 0.3–8.8 0.0171 0.015–0.020 2.708 2.632–2.783 –A 0.961 8.0SL,1 25.0TL
Chlorophthalmus agassizi ≥500 74 11.4–19.5 13.7–100.0 0.0038 0.002–0.006 3.401 3.222–3.579 +A 0.952 11.5TL,4 40.0TL
Coelorinchus caelorhincus ≥500 27 13.0–30.0 5.2–112.0 0.0006 0.0003–0.0013 3.509 3.271–3.749 +A 0.973 17.2TL,5 48.0TL
Malacocephalus occidentalis ≥500 15 27.0–38.5 49.3–162.8 0.0003 0.0002–0.0003 3.648 2.936–4.359 +A 0.904 45.0TL
Laemonema goodebeanorum ≥500 15 7.5–27.3 2.4–191.5 0.0023 0.001–0.004 3.379 3.104–3.655 +A 0.982 30.3SL
Merluccius albidus ≥500 40 27.3–40.9 212.8–699.7 0.0373 0.022–0.064 2.627 2.471–2.782 –A 0.968 23.0SL,6 70.0TL,6
Urophycis cirrata 23 20.4–43.5 86.4–770.7 0.0162 0.008–0.033 2.864 2.659–3.069 I 0.976 66.0TL
Lepophidium brevibarbe 26 11.3–28.8 4.6–117.1 0.0017 0.001–0.003 3.313 3.151–3.475 +A 0.987 10.1TL,7 27.3SL
Porichthys plectrodon 217 4.2–18.3 1.2–93.3 0.0182 0.015–0.022 2.856 2.771–2.941 –A 0.953 8.0FL,8 29.0TL
Chloroscombrus chrysurus 40 11.6–16.3 31.5–68.4 0.0182 0.017–0.018 3.000 I 0.967 11.2FL 65.0TL
Trachurus lathami 32 10.4–17.9 18.8–77.6 0.0443 0.026–0.076 2.598 2.394–2.802 –A 0.957 11.4TL 40.0TL
Citharichthys spilopterus 70 6.4–11.9 5.2–27.8 0.0283 0.021–0.038 2.763 2.632–2.894 –A 0.963 12.0SL,9 21.0TL
Cyclopsetta chittendeni 231 4.5–28.8 1.2–371.3 0.0119 0.009–0.014 3.081 3.012–3.148 I 0.972 14.5TL,9 33.0TL,9
Monolene sessilicauda 36 4.9–11.8 1.1–9.6 0.0095 0.006–0.014 2.858 2.667–3.048 I 0.964 18.0TL
Trichopsetta ventralis 873 3.6–18.0 0.5–59.6 0.0109 0.010–0.012 3.092 3.045–3.139 I 0.950 20.0TL
Symphurus diomedeanus 21 5.0–14.7 0.9–31.0 0.0067 0.004–0.012 3.169 2.927–3.411 +A 0.975 22.0TL
Trichiurus lepturus 17 7.4–65.3 0.1–103.3 0.0001 0.0001–0.0002 3.357 3.198–3.515 +A 0.993 30.0TL 234.0TL
Bembrops gobioides ≥500 21 8.8–23.4 3.9–82.6 0.0039 0.002–0.008 3.203 2.934–3.471 +A 0.970 30.0TL
Synagrops bellus 20 6.3–20.7 4.6–166.6 0.0174 0.010–0.031 3.029 2.813–3.243 I 0.979 13.0TL,13 46.0TL,14
Epigonus pandionis ≥500 56 9.8–20.2 22.8–154.2 0.0358 0.022–0.058 2.809 2.633–2.984 –A 0.950 11.2TL,15 23.5TL
Centropristis philadelphica 42 9.7–23.5 23.2–289.3 0.0323 0.020–0.053 2.862 2.676–3.047 I 0.960 30.0TL
Lutjanus campechanus 35 8.0–24.7 12.7–467.2 0.0237 0.013–0.042 3.032 2.806–3.258 I 0.958 9.41FL,11 100.0TL
Pristipomoides aquilonaris 477 3.3–20.0 1.0–197.2 0.0251 0.024–0.025 2.873 2.830–2.916 –A 0.973 56.0TL
Prionotus longispinosus 183 3.9–24.7 1.3–307.6 0.0397 0.030–0.053 2.771 2.660–2.881 –A 0.931 12.0TL,16 35.0TL
Prionotus paralatus 180 7.8–17.5 7.5–85.2 0.0142 0.011–0.018 3.056 2.959–3.153 I 0.956 10.0TL,16 18.0SL,16
Peristedion greyae 123 12.8–18.4 11.9–33.4 0.0110 0.007–0.017 2.738 2.580–2.895 –A 0.907 23.9TL
Micropogonias furnieri 26 12.0–20.2 40.4–155.5 0.0643 0.035–0.118 2.594 2.368–2.821 –A 0.959 24.3TL 60.0SL
Fowlerichthys radiosus 47 2.6–9.4 1.5–57.2 0.1357 0.105–0.176 2.578 2.411–2.744 –A 0.956 25.0TL10
Dibranchus atlanticus 178 3.4–10.8 1.5–25.7 0.0696 0.059–0.083 2.434 2.351–2.517 –A 0.957 10.9TL,17 39.4TL
Ogcocephalus declivirostris 23 6.1–10.3 6.8–37.5 0.0304 0.019–0.048 3.027 2.805–3.248 I 0.975 16.5TL
Zalieutes mcgintyi 17 3.3–7.3 1.4–10.5 0.0579 0.039–0.087 2.634 2.415–2.853 –A 0.978 10.0TL
Lagocephalus laevigatus 30 3.9–36.0 4.2–1050.3 0.0601 0.040–0.090 2.672 2.512–2.833 –A 0.976 24.5SL,12 100.0TL

We calculated the length–weight relation using the allometric formula

W = aLb

where W is the weight of the fish [g], L is the standard length [cm], a is the intercept and b is the allometric coefficient/slope. The values of a and b were calculated with Statgraphics software (Centurion XV, Version 15.1.02, Copyright 1982–2006 StatPoint, Inc.) with a linear least squares regression using a logarithmic scale. With the value of the slope (b), it was established if the fish species has negative growth (b < 3) or positive allometric growth (b > 3) and b = 3, indicating isometric growth (Froese et al. 2011). Outliers were removed using logarithmic plots, and limits for a and b were estimated by a Student’s t-test with a 95% confidence (Froese 2006). For comparison, information on the maximum length (Lmax) and the length at first maturity (Lm) is taken from FishBase and other references, with the respective length type being indexed (TL= total length, FL= Fork length). This study provides LWR that had not yet been reported for 11 species representing four different families. In some cases, when the number of specimens and/or the range of sizes was very narrow to estimate the a and b parameters of the LWR, we assumed an isometric relation (b = 3) (Froese 2006; Hay et al. 2020) and the value of the intercept (a) will be obtained with the following formula



The descriptive statistics and the estimated LWR parameters for 39 species are summarized in Table 1. All LWR estimates were statistically significant (P < 0.05), yielding R2 > 0.900. New maximum lengths are reported for four species: Cyclothone alba (5.6 cm SL), C. braueri (4.6 cm SL), C. pseudopallida (4.8 cm SL), and Lepophidium brevibarbe (28.8 SL). All the values of “a” ranged between 0.0001 (Trichiurus lepturus) and 0.1357 (Fowlerichthys radiosus); and the “b” values oscillated between 2.309 (Cyclothone alba) and 3.648 (Malacocephalus occidentalis). Positive allometric growth was reported in nine species, negative allometric growth in 16 species, and isometric growth in 14 species.

The LWR of 11 species that correspond to 10 families have not been previously reported, so it is an important contribution to their knowledge. These families and species are Congridae: Rhynchoconger flavus, Gonostomatidae: Cyclothone alba, Moridae: Laemonema goodebeanorum, Cyclopsettidae: Cyclopsetta chittendeni, Bothidae: Monolene sessilicauda, Cynoglossidae: Symphurus diomedeanus, Bembropidae: Bembrops gobioides, Triglidae: Prionotus paralatus, Antennariidae: Fowlerichthys radiosus, and Ogcocephalidae: Ogcocephalus declivirostris, Zalieutes mcgintyi.


The abundance of fish species associated with depths greater than 500 m, is usually low and the available information on their populations and growth rates are scarce (Danovaro et al. 2017). Therefore, any new data on their biology is important. The deep-sea species reported in this study are carnivorous, occurring in the vertical gradients of the continental slope and the bathyal zone, and were exemplified by Epigonus pandionis, Merluccius albidus, Chauliodus sloani, Chlorophthalmus agassizi (see Ramírez et al. 2019). Furthermore, we highlight an amplitude in its maximum length reported by the literature corresponding to Cyclothone alba from 2.9 to 5.6 cm SL, Cyclothone braueri from 3.8 to 4.6 cm SL, Cyclothone pseudopallida from 4.6 to 4.8 cm SL (Harold 2015) and Lepophidium brevibarbe from 27.3 to 28.8 cm SL. In addition, we consider that these species are the ones that are possibly being most affected during oil extraction maneuvers and hydrocarbon leaks in the depths (Fisher et al. 2016). The genus Cyclothone corresponds to the most abundant resource in these deep zones (Olivar et al. 2017) and is perhaps the main food source that generates stability in populations, so its impact would generate a disparity in the deep marine ecosystem.

LWR studies in the northern Gulf of Mexico have been very scarce. In these studies, the species analyzed include Chloroscombrus chrysurus and Citharichthys spilopterus (see Dawson 1965; Galindo-Cortés et al. 2015) and a single deep-sea species Urophycis cirrata (see Matlock et al. 1988). The majority of the species mentioned in these investigations are associated with shallow coastal areas. In the presently reported study, LWR information is provided on ecologically important species found at depths greater than 500 m, including records of both juvenile and sexually matured organisms. With this information, the reports of these species in the area were completed, as well as the delivery of new biological information on the deep-sea ecosystem, which is a poorly studied region located in the north of the Gulf of Mexico, and where samples are difficult to obtain (Blomberg and Montagna 2014). Likewise, we recorded species of Micropogonias furnieri and Citharichthys spilopterus that did not reach sexual maturity and were captured by shrimp trawls of the same dimensions as the fishing boats, so it is possible that both species are showing a decrease in their populations.

The slope (b) that was estimated in this study was between the expected range of 2.5 to 3.5 (Froese 2006), except for Cyclothone alba (2.309) and Dibranchus atlanticus (2.434) that were found below this range of values, and for Malacocephalus occidentalis which is above those values (3.648). For Cyclothone braueri and Chloroscombrus chrysurus with a low number of specimens and/or with low range sizes (Carlander 1997), the LWR was calculated assuming b = 3.0, being the value of the intercept considered by the formula of Hay et al. (2020). These low values can be attributed also to the combination of one or more of the following factors: habitat, area/season effect, gonad maturity stages, sex, stomach fullness, health condition, population, and differences within species and preservation techniques (Tesch 1971; Froese 2006; Bautista-Romero et al. 2012). Finally, a total of nine and 16 species showed positive and negative allometric growth, respectively, while isometric growth was reported in 14 species.

Author contribution

(following Contributor Roles Taxonomy of CRediT

Ariel Adriano Chi Espinola: Conceptualization, Formal Analysis, Investigation, Methodology, Visualization, Writing—original draft preparation, Writing—review and editing.

María Eugenia Vega Cendejas: Conceptualization, Funding acquisition, Investigation, Project administration, Resources, Supervision, Validation, Visualization, Writing—original draft preparation, Writing—review and editing.

Jovita Mirella Hernández de Santillana: Conceptualization, Data curation, Formal analysis, Methodology, Visualization, Writing—original draft preparation.


This is a contribution from the Gulf of México Research Consortium (CIGOM). We are grateful to Alex Acosta, María Blanqueto, Sergio Zavala and Mariana Uribe for processing the samples. This research was funded by the Mexican National Council for Science and Technology - Mexican Ministry of Energy - Hydrocarbon Fund, project 201441.


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